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The Gardens' Bulletin
Singapore
Volume 55
Singapore Botanic Gardens
2003
© Singapore Botanic Gardens
Singapore, 2003
Applications for reproduction should be
made to the Singapore Botanic Gardens,
1 Cluny Road
Singapore 259569
DATE OF PUBLICATION OF THE TWO PARTS WERE:
Part | Pp. 1-172 17 September 2003
Partt2 Pp. 173-315 15 December 2003
a
Gardens’ Bulletin Singapore 55 (2003)
CONTENTS
Chew, W.-L.
PTS. 1 Wer AOC Se eA Sere we whines aks wa dads s adaaiepumkee i iuda sccenouaabeeoanstvorsedl &
Chua, L.S.L and L.G. Saw
A new record of Dryobalanops beccarii (Dipterocarpaceae ) from Peninsular
UMMA re Net it de dao a Sete ee ee Re a piositecn us pean das vn noxteh@eunden od sar onee cag iars ntekogs sogeepaapau
Chua, L.S.L and L.G. Saw
Conservation notes on Vatica yeechongii (Dipterocarpacae) from
ae nae AOA erie Dh ou inh kus ook avon nine ules Oba08 Sevinoncpeplgnwws Dasennnasnetemnanwakasvodmeseand
Chung, R.C.K., E. Soepadmo and A.L. Lim.
The significance of pollen morphology in the taxonomy of Grewia and
Microcos (Tiliaceae) in Peninsular Malaysia and Borneo ....................0.0sss000secceeee ee 02-239
Gusman, G.
Arisaema fimbriatum (Araceae) and its intraspecific Variation ................eseeseeeeerereee ee e201
Hassan I.
Amote On ordia premnifolia Rid). (Borapinaceat) «isi. ac<cccooeetessciencdpcvadsgense so vapess ser LOD
Hay, A. and C. Herscovitch
A new species of Schismatoglottis (Araceae) from Sabah, Malaysia ...........ccceceeeeeeeees 27
Herscovitch, C. and A. Hay
An unusual new species of Homalomena (Araceae) from New Guinea ...............::ceee 31
Hilliard, O.M., B.L. Burtt and M.H. Bokhari
Pleuroschisma, a new section of Cyrtandra (Gesneriaceae) from Borneo...................0068 35
Kiew, R.
Begonia peninsulae (Begoniaceae), a confused species from Malesia ..................::00000 61
Kiew, R. and C. Geri
Begonias from the Bau limestone, Borneo, including a new SpeCi€s ..............:.::eeeeeeeeeees 113
Kiew, R. and I.M. Turner
te ity Plants CnieiIC 10 wai APOE 1 55.0 yd ned a escies dn tod dctence c dbdvcenereetenereescdaveen as Bansel 1S
Kloet, S.P. Vander
Re-examination of Vaccinium dialypetalum (Ericaceae) — Erratum ...........0:.:ccccesseeeeeees 163
Lee, S., A. Samsuri, P. Leong, Ali Ibrahim and A.-T. Gwee
A botanical survey of Chek Jawa, Pulau Ubin, Singapore ..................sscscssessscccsestee se ees 271
Mabberley, D.J.
New species of, and other notes on, Chisocheton and Walsura (Meliaceae)................... 189
Middleton, D.J.
A new species and a new combination in Bornean Kopsia (Apocynaceae:
APOCYNOIGEAREC)..........veseaceesianniesnantbuasivniiosaicncinesieh niniiov ch Geoedeeitns tual tse
Middleton, D.J.
A revision of Dyera (Apocynaceae: Rauvolfioideae) ...............csscccecccceeeeeesssreeee ee ee
Pearce, K.G.
New species and varieties of Symplocos (Symplocaceae) from Borneo...............:::0++
Pearce, K.G.
Five new Begonia species (Begoniaceae) from the Niah National Park,
Sarawak, Malaysia ..................000
Skornickova, J., M. Sabu and M.G. Prasanthkumar
A new species of Curcuma L. (Zingiberaceae) from Mizoram, India ...............eeee
Skornickova, J., M. Sabu and M.G. Prasanthkumar
Curcuma codonantha (Zingiberaceae) — a new species from the Andaman Islands,
Tinta ....ceccacaccovecscvsdeccsaeescccencsxceensnvazses<wawebuuystadaes gone ee ace ne Ese ee ee eee aaa ea
Vermeulen, J.J. and P. O’Byrne
New species and new records of Southeast Asian Bulbophyllum (Orchidaceae).........
Vermeulen, J.J. and P. O’Byrne
Six new species of Bulbophyllum (Orchidaceae) from Sulawesi ............:::cceeeeeeeeeree es
Wong, C., G. Argent, R. Kiew, Ohn Set and Y.Y. Gan
The genetic relations of Musa species from Mount Jaya, New Guinea, and
a reappraisal of the sections of Musa (Musaceacy Tittt.J2 is. .nceresscsensteoastenes en ate te
Zahid, M.S.
Three new species of Porterandia (Rubiaceae) from Mount Kinabalu, Borneo..........
Obituary
Mohamad Shah bin Hj Mohamad Noor - R.Kaiew............ 00. ...ccscccccececccceeeeee ee eeceee ones
Dr Chang Kiaw Lan - K.M WOmtng.......020co: oe0ss0% «050 s00nse oysodnvsxsntranaaeen ian
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Singapo
THE GARDENS’ BULLETIN
The Garden’s Bulletin Singapore publishes papers on plant taxonomy (including
revisions), horticulture, phytogeography, floristics, morphology, anatomy and related
fields with emphasis on plants in the West Malesian region.
EDITORIAL BOARD
Dr Ruth Kiew
(Editor)
Singapore Botanic Gardens
Dr P.Y. Tan
(Assist. Editor)
Singapore Botanic Gardens
Dr S. C Chin
Singapore Botanic Gardens
Dr M.J.E. Coode
Royal Botanic Gardens
Kew, U.K.
Dr R.T. Corlett
University of Hong Kong
Hong Kong
Dr M.C. Roos
National Herbarium of Netherlands
Leiden University Branch, Netherlands
Dr E. Soepadmo
Forest Research Institute Malaysia
Kepong, Malaysia
Dr W.K. Tan
Singapore Botanic Gardens
The Gardens’ Bulletin is published twice yearly by the National Parks Board,
Singapore. Neither the National Parks Board not the Editorial Board is responsible
for the opinions or conclusions expressed by the contributing authors.
The annual subscription for the Gardens’ Bulletin is Singapore $100.00
including postage. Overseas subscribers are required to make payment in the form of
bank drafts or international money orders in Singapore currency payable to National
Parks Board, Singapore.
Instructions for contribiting authors are found on the inside backcover.
‘ * 0 3 2003 ‘ et acl
The Gardens’ Bulletin
Singapore
VOL. 55 (Part 1) July 2003 ISSN 0374-7859
CONTENTS
Chua, L.S.L. and L.G. Saw
A new record of Dryobalanops beccarii (Dipterocarpaceae) from
Fane VSB Bs Sete fas ees tetrad! oc aes:! use 1, oily. dgee iy als CMa aweeeseceoe d
Chua, L.S.L. and L.G. Saw
Conservation notes on Vatica yeechongii (Dipterocarpaceae) from
Be eRe Nh Nts os os dace pee damnns weg dees give ndeeeoebiod bani es ‘4
Chew, W.-L.
SLMS th WIRMeSIal PigeraCege Beis tihs anys vein ches iencs'ssl bite qepehgemedovenxdeeoveheers.y0004 3
Hay, A. and C. Herscovitch
A new species of Schismatoglottis (Araceae) from Sabah, Malaysia ............... 27
Herscovitch, C. and A. Hay
An unusual new species of Homalomena (Araceae) from New
aN a BS a cn ae cctas as atid aanicd apuumpt nc cksen ipiscsie davcpibaiine 31
Hilliard, O.M., B.L. Burtt and M.H. Bokhari
Pleuroschisma, a new section of Cyrtandra (Gesneriaceae) from
EE re TURE Node tees et Ae ee A EP =
Kiew, R.
Begonia peninsulae (Begoniaceae), a confused species from Malesia..................04. 61
Middleton, D.J.
A new species and a new combination in Bornean Kopsia
AR ON NEN IN Fetes Niet nso a ia lsvannepecigeraneenasesseusvencasdncwnnese 65
Pearce, K.G.
New species and varieties of Symplocos (Symplocaceae) from Borneo................ 69
Pearce, K.G.
Five new Begonia species (Begoniaceae) from the Niah National Park,
Sarawak, Malaysia... sisccccck cited escarole Ste a 73
Skornickova, J., M. Sabu and M.G. Prasanthkumar
A new species of Curcuma L. (Zingiberaceae) from Mizoram, India.................... 89
Wong, C., G. Argent, R. Kiew, Ohn Set, and Y.Y. Gan
The genetic relations of Musa species from Mount Jaya, New Guinea,
and a reappraisal of the sections of Musa (Musaceae) .............cscssecccsesseeeee eee iD]
Kiew, R. and C. Geri
Begonias from the Bau Limestone, Borneo, including a new species ................. 113
Vermeulen, J.J. and P. O’Byrne
New species and new records of Southeast Asian Bulbophyllum
(Orchidaceae) -.2.2i0c ssc csceceswsieattderstarresliecevave fess pest Le ee 125
Kloet, S.P. Vander
Re-examination of Vaccinium dialypetalum (Ericaceae) — Erratum ................... 163
Obituary
Mohamad Shah bin Hj Mohamad Noor ................. 00. seessseeeeeeeeeeeee RY Kiiew-165
Date of Publication: 17 September 2003
Published by
National Parks Board
Singapore Botanic Gardens
1 Cluny Road
Singapore 259569
Printed by
Print Dynamics (S) Pte Ltd
Garden’s Bulletin Singapore 55 (2003) 1-6
A New Record of Dryobalanops beccarii (Dipterocarpaceae)
from Peninsular Malaysia
L.S.L. CHUA AND L.G. SAW
Forest Research Institute Malaysia,
Kepong 52109, Kuala Lumpur, Malaysia
Abstract
Dryobalanops beccarii Dyer, Dipterocarpaceae, first described and recorded from Borneo,
has now been documented from Johore, Peninsular Malaysia, where it grows on ridges in
several compartments in the Panti Forest Reserve, Kluang Forest Reserve, and the Labis Forest
Reserve. It is a large tree frequently reaching 40 m height and more than 1 m diameter.
Introduction
Two species of Dryobalanops, D. aromatica and D. oblongifolia, were recorded for
Peninsular Malaysia (Ashton, 1982). The genus Dryobalanops is clearly distinguished
from other dipterocarp genera by the presence of five wing-shaped, subequal fruit
calyx lobes and closely parallel venation on its leaf blade. In Peninsular Malaysia,
trees of both species are characterized in the field by scaly bark, peeling in large,
irregular, thin flakes. In D. oblongifolia, leaves are oblong or oblong-lanceolate and
the fruit calyx lobes are shorter than the nut, while in D. aromatica, leaves are oval or
sub-ovate and the calyx lobes are longer.
During a plant diversity enumeration exercise in the Virgin Jungle Reserves
(VJR) network in Peninsular Malaysia, Ang Khoon Cheng and Kamarudin Saleh
discovered in Kluang Forest Reserve (FR) and the VJR in Labis FR, Johore, trees
that had leaves that closely resembled D. aromatica but the bark was distinctly different
in that the outer bark was scaly, very shallowly fissured and had a laminate inner
bark. It was clear that these populations belonged neither to D. aromatica nor D.
oblongifolia but their identity could not be confirmed in the absence of flowers and
fruits.
A dipterocarp mast-fruiting episode occurred in Peninsular Malaysia during
the second and third quarters of 2002. Taking advantage of this cyclic event, teams
from the Botany and Seed Technology Units, Forest Research Institute Malaysia
(FRIM), launched an extensive fruit collection exercise, targeting species that are
rare and/or threatened. Fruiting trees in several of these Dryobalanops populations
i)
Gard. Bull. Singapore 55 (2003)
enabled materials for herbarium specimens and fruits for ex situ planting to be
collected. The identity of the species could be confirmed and proved to be D. beccarii
Dyer. A full description of the species based on characters from these populations is
provided below.
Dryobalanops beccarii Dyer J. Bot. 12 (1874) 100; Ashton, Fl. Malesiana I (9)
(1982) 375. Type: Malaysia, Sarawak, Matang, Beccari PB2944 (K, lecto).
Large tree to 30 m tall, 63.7 cm diam., with small buttress to 1.5 m tall. Bark shallowly
fissured to scaly, flaky and peeling off in pieces on the lowest part of the trunk
immediately above the buttress, greyish-white; inner bark laminate, reddish-brown;
sapwood yellowish brown, exuding resin when cut. Twigs 3 mm diam., glabrous,
drying almost black. Leaves thickly coriaceous, glabrous, drying brown above, dark
brown below; petiole 10-17 mm long, 1 mm thick, glabrous, drying almost black;
blade broadly ovate, 3.5—6.3 x 2.4—3.7 cm, apex tapering to a 7-mm long acumen,
base broadly cuneate to rounded, margin thickened, midrib prominently raised below,
sunken above, secondary veins numerous, closely parallel, faint on both surfaces,
smelling of camphor smell when crushed. Flowers not seen. Fruit calyx glabrous,
shallow cup tapering gradually and cuneate at the pedicel, lobes 5, equal, oblong-
spathulate, base to 2 mm, yellowish green when fresh, 4.0-6.5 x 0.6—1.0 cm, veins
conspicuously raised on both surfaces, nut ovoid, glabrous, yellowish green when
fresh, to 1.4 x 0.9 cm, style remnant 1 mm long, free from calyx except the base.
Vegetatively, the leaf morphology of Dryobalanops aromatica and D. beccarii
is very similar and hence it is difficult to differentiate them when sterile. However,
the bole character as described above can readily distinguish the two species. In
addition, the fruits of D. beccarii are different from D. aromatica (Table 1).
Table 1: The bark and fruit characters of Dryobalanops aromatica and D. beccarii
D. aromatica D. beccarii
Outer bark long, thin, recurved, irregular shallowly fissured to scaly
scales
Inner bark fibrous laminate
Fruit calyx size (cm) 4.0-6.0 x 0.8-2 4.0-6.5 x 0.6—1.0
Max. nut size (cm) 3% 1 14x09
Specimens examined: Johore: Compartment 6B in Panti FR, Damahuri S. FRI 43567,
7 August 2002 (KEP), Ang K.C. FRI 43651, 13 August 2002 (KEP); Kluang FR, Ang
K.C. FRI 43653, 14 August 2002 (KEP).
Dryobalanops beccarii in Peninsular Malaysia 3
Geographical distribution in Peninsular Malaysia
In Peninsular Malaysia, Dryobalanops beccarii is known only from Johore from three
localities, namely in compartments 6A, 6B, 7, 8A and 8B, Panti FR; in compartment
132, Kluang FR; and in compartment 841 in VJR, Labis FR.
Prior to this discovery, D. beccarii was only known to occur in Borneo on
leached sandy soils on coastal hills and inland ridges below 700 m (Ashton, 1982).
Like D. aromatica and D. oblongifolia, it shows the same pattern of geographic
distribution in southern Johore and west Borneo. Corner (1958) has drawn attention
to the similarity in the floras of these two regions, which he termed the Riouw pocket.
Ecology
In Johore, this species is found in the lowland dipterocarp forest at about 75 m altitude,
confined to ridges. They are large trees in the emergent and main canopies. On some
ridges, it is co-dominant with Shorea curtisii Dyer ex King. In Kluang FR, it grows
sympatrically with D. aromatica on steep hill slopes. In all the three sites, D. beccarii
grows gregariously. In Borneo, the species is locally abundant on leached sandy soils
on coastal hills and inland ridges below 700 m.
Germination
Three seed batches of mature fruits were collected from five trees in Panti FR and six
trees in Kluang FR in August 2002 and germinated with the seed wings removed in
the FRIM nursery (Table 2). Germination is deemed to begin when the radicle protrudes
through the seed coat. Germination is epigeal. The hypocotyl elongates to c. 1.8 cm.
The emergent cotyledons are bilobed, fleshy and unequal. The first two leaves are
opposite. In some seedlings, branching may begin at the first node.
Mean percentage germination was 73 + 12 s.d., the percentage varying from
62 to 86 (Table 1). All three seed batches showed a typical germination sigmoid curve
(Fig. 1).
The germination period for Dryobalanops beccarii seed batches was longer
than that of D. aromatica (100% within 20 days, Ng 1991, Siti Asha er al. 1995,
Tamari 1976) and D. oblongifolia (100% within 10 days, Siti Asha et al. 1995 and 33
days, Ng 1991). In addition to the longer germination period, the three seed batches
of D. beccarii achieved a maximum germination percentage of only 86.
Conservation notes
In Panti FR, ground checks in August 2002 indicated that the boundary of
compartments 6B, 8A and 8B, which forms part of the forest reserve boundary, is
adjacent to land being developed for small-scale agriculture activities. Because
agriculture has a propensity for expansion, it is recommended that the forest office
regularly monitors this section of the boundary to ensure its integrity and prevent
future conflict. Compartments 6B, 8A and 8B had been logged in 1982/83. These
4 Gard. Bull. Singapore 55 (2003)
compartments are neither VJR nor are they above 1000 m elevation and hence have
no legal protected status and may be licensed out for timber harvesting. Populations
in Labis VJR and in Compartment 132, Kluang FR have a much brighter prospect as
these forests are virgin and VJR and water catchment areas have protected status.
Studies are currently being undertaken to determine the species’ population size and
genetic structure.
In view of the above scenario, Dryobalanops beccarii in Peninsular Malaysia
is given the 2001 IUCN category of Endangered EN A3dB1. D. beccarii falls within
the A3 category (there will be a projected or suspected population size reduction of
250% within the next 10 years or three generations, whichever is the longer);’d’ (the
reduction can be based on actual or potential levels of exploitation); and B1 (its
geographic range, in the form of the extent of occurrence, is estimated to be less than
5000 km’).
Table 2. Percentage germination and survival in three seed batches of Dryobalanops
beccarii
Batch No. 2002-0502 (FRI 43567) 2002-0545 (FRI 43653) FRI 43651
Locality Panti FR, Compartment Kluang FR Panti FR,
6B Compartment
6B
Collector Damahuri S Ang KC Ang KC
Date planted 12 August 2002 16 August 2002 16 August
2002
No. sown 4] 617 78
Max. germination (%) 71 86 62
Germination period 4—32 4-3] 2-31
(days)
No. days to achieve 6 9 23
50 % germination
% seedling survival 22.8 72.8 100
after 1 month
Dryobalanops beccarii in Peninsular Malaysia 5
100
Germination (%)
Nn
S
Days
—e— 2()02-0502 —m— 2002-0545 —&— FRI 43651
Figure 1: Germination curve for three different seed
batches of Dryobalanops beccarii
Acknowledgements
We thank Hj. Ros and the foresters and rangers of the South Johor District Forest
Office and Hj. Ahmad Fekri and staff of the Kluang District Forest Office for the
permission and field assistance provided; Peter S. Ashton for confirming species
identity; Kamarudin Saleh and Ang Khoon Cheng (FRIM) who first noted the presence
of Dryobalanops beccarii and together with Damahuri Sabari, Mohd. Aidil Nordin,
Mustapa Data and Ayau Kanik collected sufficient data for confirmation. Financial
support from the Flora Malaysiana Centre Fund, Ministry of Primary Industries,
Malaysia under Project 6: Conservation Monitoring System for Threatened Plants is
gratefully acknowledged.
6 Gard. Bull. Singapore 55 (2003)
References
Ashton, P.S. 1982. Dipterocarpaceae. Flora Malesiana. Ser. 1, 9: 237-552.
Corner, E.J.H. 1958. An introduction to the distribution of Ficus. Reinwardtia 4:
15-45.
IUCN, 2001. JUCN Red List Categories. Version 3.1. 1UCN Species Survival
Commission. IUCN, Gland, Switzerland and Cambridge, UK.
Ng, F.S.P. 1991. Manual of Forest Fruits, Seeds and Seedlings. Malayan Forest
Records No. 34, Vol. 1. Forest Research Institute Malaysia. Kuala Lumpur. 400
PP.
Siti Asha A.B., K.C. Ang, and S. Zaiton. 1995. Records of germination trials of
dipterocarp seeds at FRIM, Kepong, 1979-1994. Research Data No. 4. Forest
Research Institute Malaysia. Kuala Lumpur. 161 pp.
Tamari, C. 1976. Phenology and seed storage trials of Dipterocarps. FRI Research
Pamphlet No. 69. Forest Research Institute Malaysia. Kuala Lumpur. 73 pp.
Garden’s Bulletin Singapore 55 (2003) 7-12
Conservation Notes on Vatica yeechongii (Dipterocarpaceae)
from Peninsular Malaysia
L.S.L. CHUA AND L.G. SAW
Forest Research Institute Malaysia
Kepong 52109, Kuala Lumpur, Malaysia
Abstract
The conservation status of Vatica yeechongii L.G. Saw, recently discovered from Peninsular
Malaysia, is described.
Introduction
Vatica yeechongii was discovered along Sungai Tekala in Sungai Tekala Recreational
Forest, which is located in Sungai Lalang Forest Reserve, Selangor, Peninsular
Malaysia, 3° 03.485’ N, 101° 52.373’ E, alt. 79 m asl (Saw, 2002). Several weeks
after its discovery in the type locality, a population was found in Compartment 4,
Setul Forest Reserve, along the upper stream of Setul River, Negeri Sembilan
(2°46.937’ N, 101°55.069’ E). The population was located in the forest margin of a
logged-over forest adjacent to the trunk road from Mantin to Seremban town at an
altitude of c. 192 m.
Vatica yeechongii is very distinct in leaf characters and nut shape (Saw, 2002).
The leaf blade is oblanceolate, 44-84 x 10—6.5 cm, thickly coriaceous, bullate above
with 28—30 pairs of veins. The nut is ovoid.
Ecology
A medium-sized tree 8—15 m tall and 9-13 cm diameter, V. yeechongii occupies the
dense understorey of lowland dipterocarp forest. In the type locality, nine trees were
found on gentle earth banks of a free flowing stream near the public campsite. There
were no trees further upstream. Details of its growth pattern are described in Saw
(2002). Large dipterocarps such as Shorea leprosula Miq., S. parvifolia Dyer, S.
macroptera Dyer, S. acuminata Dyer, S. pauciflora King, S. ovalis (Korth.) Blume,
S. bracteolata Dyer, S. dasyphylla Foxw., Dipterocarpus crinitus Dyer and D.
cornutus Dyer dominated the forest. Non-dipterocarps included /ntsia palembanica
4 Gard. Bull. Singapore 55 (2003)
Mig. (Leguminosae), Saraca cauliflora Baker (Leguminosae), Terminalia
subspathulata King (Combretaceae), Pometia pinnata J.R. Forst. & G. Forst.
(Sapindaceae), Canarium patentinervium Mig. (Burseraceae), Elateriospermum tapos
Blume (Euphorbiaceae), Palaquium gutta (Hook.f.) Baill. (Sapotaceae),
Pimelodendron griffithianum (Miull.Arg.) Benth. (Euphorbiaceae), Endospermum
diadenum (Miq.) Airy Shaw (Euphorbiaceae), Dyera costulata (Miq.) Hook.f.
(Apocynaceae), Artocarpus lanceifolius Roxb. (Moraceae), A. scortechinii King,
Gironniera subaequalis Planch. (Ulmaceae), G. nervosa Planch., Payena lucida A.
DC. (Sapotaceae), Streblus elongatus (Miq.) Corner (Moraceae), Litsea costalis (Nees)
Kosterm. (Lauraceae), Gynotroches axillaris Blume (Rhizophoraceae), Cratoxylum
formosum (Jack) Dyer (Guttiferae), Scaphium macropodum (Miq.) Beumée ex Heyne
(Sterculiaceae) and Pternandra echinata Jack (Melastomataceae).
At Setul, the forest margin vegetation gradually gave way to a forest having
a main canopy to c. 20 m tall, dominated by Shorea leprosula, S. macroptera, S.
multiflora (Burck) Symington, S. acuminata, §. parvifolia, Parkia speciosa Hassk.
(Leguminosae), Pometia pinnata, Campnosperma auriculatum (Blume) Hook.f.
(Anacardiaceae), Artocarpus scortechinii, Pterocymbium tinctorium (Blanco) Merr.
(Sterculiaceae) and Alstonia angustiloba Miq. (Apocynaceae). The understorey lacked
palms and climbing rattans and the forest floor was covered with abundant dipterocarp
seedlings. Herbs such as Tacca integrifolia Ker Gawl. (Taccaceae) and Costus speciosus
(J. KOnig) Sm. (Costaceae) were present in scattered numbers. At least 70 V. yeechongii
trees were found on steep slopes of riverbanks and on slopes away from the riverbanks.
Trees here averaged 9.8 m tall (range 2—21 m), had a diameter at breast height c. 8.2
cm (range 1.6—18.8 cm) with a spreading crown. Its bark was whitish and smooth
with horizontal rings. Those growing along the riverbanks had poor bole form and
multiple branching was common while those further away had a better bole form.
Germination
Mature fruits were available from at least six trees in Sungai Tekala and one tree in
Setul Forest Reserve from June to August 2002. Three seed batches, collected from
two localities at different dates, were germinated with the seed wings removed and
observed in the Forest Research Institute Malaysia, FRIM (Table 1). Germination is
deemed to begin when the radicle protrudes through the seed coat. Germination of
Vatica yeechongii is epigeal. The emergent cotyledons are bilobed, fleshy and equal.
The hypocotyl elongates to c. 4.8 cm tall and the first two leaves are opposite.
Mean percentage germination was 63.011 +31.207 s.d., the percentage varying
from 42.1 to 98.9. All three seed batches showed a typical sigmoidal germination
curve. Germination percentage on the first day of germination, i.e. day 5, was 38%
for seed batch 2002-0510 and 18.7% for seed batch 2002-0503. Despite this promising
Vatica yeechongii 9
Table 1. Percentage germination and survival of three seed batches of Vatica
yeechongii
Batch No. 2002-0503 (FRI 46613) 2002-0510 2002-0542 (FRI 46668)
Locality Sungai Tekala, Sungai Sungai Tekala, Setul Forest Reserve,
Lalang Forest Reserve, Sungai Lalang Negeri Sembilan
Selangor Forest Reserve
Collector Chung R Saw LG Chan YC & Ayau K
Date planted 14 July 2002 28 July 2002 20 August 2002
No. sown 107 fie, 89
Max. germination (%) 42.1 48.1 98.9
Germination period 5-37 5-25 2-25
(days)
No. days to achieve 45 32 2
40% germination
% seedling survival 5.9 97.3 40
after 1 month
beginning, however, these two seed batches took a longer time to achieve the 40%
germination mark compared to seed batch 2002-0542 (Fig. 1). Preliminary germination
results suggest that seed viability is higher in the population at Setul Forest Reserve
but this could be due to a variety of factors e.g., fruit maturity, seed vigour, timing of
fruit collection, tree health, pest infestation on fruits, etc. In addition, population sizes
could also affect gene flow; a larger population is probably more genetically diverse.
In situ Conservation
Vatica yeechongii is so far known only from these two localities. Preliminary
observations on population size suggest that this species is a rare endemic. The
population at Sungai Tekala lies in recreational forest. Under the National Forestry
10 Gard. Bull. Singapore 55 (2003)
100
80
S60
=
—
= 40
=
5
Oo 2
0
12.3.4. 5 6.7, 8,9 1011 12.13)14 15.16.17, 18,19-98 2128
Days
—o— 20()2-0542 —a— 2002-0503 —a— 2002-0510
Figure 1: Cumulative germination percentage for
different seed batches of Vatica yeechongii
Act, no harvesting of resources is permitted in such forests as they are strictly meant
for recreation. Population loss will only take place if the Forest Reserve or parts of
the reserve, through de-gazettement, are converted to non-forestry land use. For Sungai
Tekala, such action is, however, highly unlikely as the area is in the vicinity of the
water catchment for the Semenyih Dam, a principal dam providing water supply to
Kuala Lumpur. In terms of man-made threats to the reduction in population size and
area of occupancy, there are no immediate foreseeable pressures either from habitat
loss or degradation or timber harvesting. We note that the practice of clearing
undergrowth in areas designated for recreational use may affect regeneration and
survival of populations. Careful observa tions on the individuals here suggest that the
species has no potential commercial value, as trees do not attain significant harvestable
size at maturity and has a poor bole form. The Forest Department of Central Selangor
District has been alerted to the presence of the new species and steps are being taken
to conserve the population.
The population at Setul faces a different scenario. Although it lies in the
forest reserve, it is adjacent to the main road. It has some form of protection by virtue
of it being in Forest Reserve, but the fact that it lies very close to the road could pose
Vatica yeechongii ‘il
future conservation conflicts when road expansion is required. It is therefore
recommended that more stringent protection measures be provided for this
population. In view of the above scenario, Vatica yeechongii should be given the 2001
IUCN category of Critically Endangered CR A2cB1. It falls within the A2 category
(inferred or suspected population size reduction of >80% over the last 10 years or
three generations, whichever is longer); ‘c’ (a decline in area of occupancy, extent of
occurrence and/or quality of habitat) and B1 (its geographic range, in the form of the
extent of occurrence and area of occupancy, is estimated to be less than 100 km?). An
attempt to estimate the population size of the species in each locality is currently
being undertaken.
From the viewpoint of conservation, the discovery of a new dipterocarp species
in Selangor and Negeri Sembilan is remarkable because in comparison to other states,
these two states encounter higher conflict between socio-economic development and
conservation. Both populations face an acute need for non-forestry related land
development and hence there exist tremendous pressures leading to habitat loss and
degradation. The fact that a new species was discovered in this current era of
diminishing forest areas merely shows that we do not know enough about our plant
diversity and that long-term field observations are necessary. This can only take place
with concerted botanical research programmes having full national, financial and
personnel support.
Ex situ Conservation
One month after potting, 215 seedlings from the three batches survived, each bearing
at least two pairs of leaves. Distribution to other gardens and arboreta is envisaged in
the near future.
Acknowledgements
We thank Azid Adam, Mohd. Jantan, Borhan Mat Saad and Abu Kasim Omar, Selangor
District Forest Office, Cheras, Kuala Lumpur and Hj. Mohamad Hj. Ismail of the
Negeri Sembilan Barat District Forest Office for the permission and field assistance
provided; Chan Yee Chong who discovered the species; Damahuri Sabari, Mohd.
Aidil Nordin, Markandan Moorthy and the support staff of Botany Unit, FRIM, for
the field collection and ex situ maintenance of plants. Financial support from the
Flora Malaysiana Centre Fund, Ministry of Primary Industries, Malaysia under Project
6: Conservation Monitoring System for Threatened Plants is gratefully acknowledged.
12 Gard. Bull. Singapore 55 (2003)
References
IUCN, 2001. JUCN Red List Categories. Version 3.1. IUCN Species Survival
Commission. IUCN, Gland, Switzerland and Cambridge, UK.
Saw, L.G. 2002. A new species of Vatica (Dipterocarpaceae) from Peninsular Malaysia.
Gardens’ Bulletin Singapore. 54: 247-251.
Garden's Bulletin Singapore 55 (2003) 13-25
Studies in Malesian Piperaceae 3!
WEE-LEK CHEW
12 Robinson Street
Chatswood NSW Australia
Abstract
Thirteen species, mostly from New Guinea, are dealt with in this paper, of which 12 other
species previously considered distinct are reduced to synonymy. No new taxon is proposed.
Introduction
This is part of a series of papers, albeit variously titled, resulting from continuing
work initiated some 35 years ago. As mentioned previously (vide Chew, Journal Arnold
Arboretum 53 (1972) 1-25), the work began by way of an undertaking to identify the
collection of Piper made by the Royal Society Expedition to the Solomon Islands in
1965. This necessitated reference to the New Guinea species of Piper which then
extended to an assessment of the taxonomic status of the more than 150 species
described by Casimir de Candolle from that region and surrounding islands.
I was encouraged to this much larger task by the fact that the type materials
from the former German New Guinea held in the Berlin Herbarium are still extant,
having been removed to safety prior to the destruction of the Herbarium, and assisted
further by the availability of a vast collection of high quality specimens made since
about 1945.
Thus, I was enabled to determine (at the least to assess, in a few cases) the
taxonomic status of some 108 of de Candolle’s species and to identify over 1,000
collections from New Guinea and the surrounding region. This resulted in the reduction
of 75 Candollean names to synonyms. The assignment to synonymy of this large
number of de Candolle’s species should not detract from the otherwise excellent work
accomplished by this great taxonomist: he did not have the advantage of the availability
of the vast range of contemporary collections that I have.
Apart from the floristic account of the alpine species by van Royen in The
Alpine Flora of New Guinea 3 (1982) 1267-1287, I am not aware of any major study
having been undertaken on the New Guinea species since commencement of my
work.
‘Following: Blumea 20 (1972) 145-149 and Blumea 37 (1992) 159-164.
14 Gard. Bull. Singapore 55 (2003)
There still remains quite a number of Candollean species requiring
determination of their taxonomic status. Additionally, large quantities of unnamed
materials in many herbaria await identification, some of which may represent
undescribed species. As this is a large undertaking, it may need the assistance of
future botanists to continue the work.
Unless otherwise stated, all collections cited in this paper have been seen
and studied by me personally.
1. Piper abbreviatum Opiz in Presl., Rel. Haenk. 1 (1828) 157; Quisumbing, Philip.
J. Sci. 43.1 (1930) 58, fig. 24 & 25; Chew, J. Arn. Arb. 53 (1972) 1. Type: Philippines,
Luzon: Haenke s.n. (holotype PR).
Synonyms: Piper minus K. Schum. & Lauterb., Fl. Schutzgeb., (1900) 258, syn. nov.
Type: New Guinea, Ramufluss, Lauterbach 2611 (holotype B).
Piper subnudilimbum C.DC., Bot. Jahrb. 55 (1918) 205, syn. nov. Type: New Guinea,
Djamu, Schlechter 17347 (holotype B, iso K).
Piper quintuplinervium C.DC., Candollea, 1 (1923) 203, 204; Candollea, 2 (1925)
224 syn. nov. Type: New Guinea, sine loc., Lauterbach 605 (isotype WRSL).
Dioecious glabrous climber. Lamina shortly petiolate, elliptic, ovate to broadly ovate,
c. 10 x 5 cm, length:breadth ratio c. 2:1, glabrous on both surfaces, apex acute to
acuminate, base cuneate to rounded, symmetrical; lateral veins 4 pairs, the lowermost
pair very short, arising from the base, the 2nd and 3rd usually long, arising from the
midrib a little above the base, the distal 4th pair very short, often absent, arising from
the midrib near the apex. Stipules to 1.5 cm long, often 1 cm long, usually as long as
petioles. /nflorescences shorter than leaves, peduncle c. 1.5 cm long, usually as long
as petioles; males thin, to 6 cm long, bracts orbicular, peltate, subsessile; females
shorter and thicker, c. 2 x 0.8 cm, bracts orbicular, peltate, sessile. Male flowers
2-staminate; pedicel very short stout, hirsute; stamens c. 1 mm long, anthers reniform
to subglobose, 2-valved, slightly shorter than filaments. Female flowers sessile;
stigmas 3- or 4-fid, sessile. Fruits sessile, entirely concrescent at maturity.
Distribution: Philippines, Java, Sulawesi, Maluku, New Guinea, Bismark Archipelago
and Solomon Islands.
Notes: Piper minus is a smaller-leafed version of P. abbreviatum and there are no
essential differences that can justify their being kept apart. Kajewski 2047 has leaves
that vary greatly from the narrowly ovate c. 4 x 1.5 cm to broadly ovate c. 6 x 4 cm.
In Piper subnudilimbum, the type materials contain immature inflorescences.
The leaves are large, much like those of the Philippine populations of P. abbreviatum,
but the leaf bases are like the typical New Guinea plants with three pairs of lateral
Malesian Piperaceae 3 15
veins. This difference is, however, not significant and cannot be relied on to distinguish
P. subnudilimbum from P. abbreviatum.
As for Piper quintuplinervium, the isotype shows two leaf forms: the narrowly
ovate, small ones borne on end of free branches and the large pentagonal leaves borne
at the nodes of the climbing parts of the plant. Other than small differences between
P. quintuplinervium and P. abbreviatum, there is nothing significant to keep the two
species apart.
Specimens examined: NEW GUINEA: Djamu Distr., Schlechter 17347; Ramufluss,
Lauterbach 2611; sine loc., Lauterbach 605. SOLOMON ISLANDS: Kajewski 2047
(A).
2. Piper arfakianum C.DC. in Gibbs, Phyt. Fl. New Guinea. (1917) 127. Type: New
Guinea, [rian Jaya: Arfak Mts., Angi Lake, L.S. Gibbs 5525 (isotypes K, L).
Synonym: Piper pilosulinodum C. DC. in Gibbs, Phyt. Fl. New Guinea. (1917) 128,
syn. nov. Type: New Guinea, Arfak Mts., Koebre Ridge. L.S. Gibbs 5624 (holotype
BM; isotype K).
Climber, usually glabrous, rarely hairy. Leaves with 1—2 cm-long petioles. Lamina
symmetrically ovate, c. 9 x 4 cm; lateral veins 3 pairs, arising from the base; apex
long acuminate; base rounded, not auriculate. /nflorescence longer than leaves;
peduncle longer than petiole. Bracts peltate, circular. Flowers sessile, crowded. Stigma
3-fid, reflexed, subsessile.
Distribution: New Guinea.
Notes: Piper pilosulinodum, also from the Arfak Mountains, is conspecific with P.
arfakianum. Having compared the type materials of these two species and with other
recent collections, | am unable to keep the species apart. Of the collections cited
above, Kanehira & Hatusima 13708 is the closest match to the type material of P.
arfakianum and van Royen & Sleumer 7456 is a good match to the type material of P.
pilosulinodum.
Piper arfakianum is closely related to P. macropiper Pennant from which it
can be distinguished by the above enumerated combination of characters, especially
the petiole being much longer than in the latter species and the lamina base not
auriculate.
Specimens examined: IRIAN JAYA: Arfak Mt., alt. 2200 m, Kanehira & Hatusima
13708, April (A, BO); alt. 2400 m, L.S. Gibbs 5525 (BM, K); Koebre Ridge, L.S.
Gibbs 5624 (BM, K); Mt. Nettoti, alt. 1920 m, van Royen & Sleumer 7456, Nov (A,
16 Gard. Bull. Singapore 55 (2003)
BRI, CANB, L, LAE). PAPUA NEW GUINEA: Central Division, Mt. Tafa, alt.
2300 m, Brass 4047 (BRI), 5029 (BRI), 5/76 (BRI); Chimbu Distr., Mt. Wilhelm,
alt. 2600 m, Borgmann 292, Aug. (LAE); Milne Bay Distr., Mt. Dayman, alt. 2200
m., Brass 22534, May (A, CANB, LAE); Morobe Distr., Edie Creek, Ridsdale NGF
30266, Nov (LAE); alt. 2000 m, Womersley & Thorne NGF 12837, Jun. (BRI, CANB,
LAE); Mt. Kaindi, alt. 2200 m, Brass 29732, May (LAE); alt. 2438 m: Coode &
Katik NGF 32863, Jun. (BRI, CANB, LAE); alt. 2300 m, Millar NGF 23627, Aug.
(BRI, CANB, LAE); alt. 2100 m, Streimann NGF 30867, Aug (BRI, LAE, NSW);
Southern Highland Distr., Anga, alt. 2000 m, Schodde 1556, Jul. (A, CANB, LAE);
Spreader Divide, alt. 2255 m, Streimann & Kairo NGF 45446, Nov (CANB, NSW).
3. Piper austro-caledonicum C.DC. in DC. Prodr. 16.1 (1869) 346; Schlechter,
Bot. Jahrb. 39 (1906) 92: Moore, J. Linn. Soc. Bot. 45 (1921) 381; Guillaumin,
Flore analytique et synoptique de la Nouvelle-Caledonie, Phanerogames. (1948)
90. Type: sine loc., Forster s.n. (syntype: BM). sine loc., Vieillard 1227 (syntype:
P).
Synonyms: Piper paitense Schltr., Bot. Jahrb. 39 (1906) 92, syn. nov. Type: New
Caledonia, Mt Paita. Schlechter 14964 (isotypes L, P, WRSL).
Piper comptonii S.Moore, J. Linn. Soc. Bot. 45 (1921) 381, syn. nov. Type: New
Caledonia, Mt Panie. Compton 1805 (isotype BM).
Piper peekelii C.DC., Bot. Jahrb. 57 (1922) 354, syn nov. Type: Neu-Mecklenburg,
Namatanai, P. Peekel 322 (holotype B).
Glabrous climber. Leaves petiolate. Lamina of lower leaves broadly ovate to somewhat
rounded to cordate, c. 9 x 7 cm, those of the upper leaves usually smaller; apex
shortly blunt-acuminate; base rounded to peltate, slightly unequal, glabrous on both
surfaces. /nflorescences slender, as long as the leaves, peduncle as long as the petiole:
bracts pedicellate. Male flowers 3-staminate, stamens subsessile. Female flowers
with 3-fid stigmas.
Distribution: New Guinea, Bismarck Archipelago, Solomon Islands and New
Caledonia.
Notes: Forster f. misidentified his collection from New Caledonia as Piper siriboa
Forst. f. (auct. non L.). I have seen both Forster’s and Vieillard’s collections annotated
by C.DC. and am confident that the collections belong to P. austro-caledonicum.
Based on examination of the type specimens, P. paitense, P. comptonii and P. peekelii
are all conspecific with this species because I cannot differentiate their type specimens
from specimens of P. austro-caledonicum. They are therefore reduced to synonymy.
Piper austro-caledonicum is closely related to P. elbertii C.DC. of Indonesia and P.
Malesian Piperaceae 3 17
retrofractum Vahl of the Philippines. Further study is needed to assess their relationship.
Specimens examined: NEW GUINEA: Torricelli, Schlechter 14381] (BO).
BISMARCK ARCHIPELAGO: New Ireland (Neu-Mecklenburg), Namatanai, P.
Peekel 322 (B). SOLOMON ISLANDS: Guadalcanal, Wanderer Bay, BS/P 12247
(LAE); Kolombangara, Kokove area, BS/P 7579 (LAE); New Georgia, Kibukibu River,
BSIP 4819 (LAE); Ranongga Island, BS/P 15572 (LAE); Santa Ysabel, Jejevo River,
BSIP 4700 (LAE), Sigana, Brass 3451 (A, BRI), Vuranimala village, BSJP 1540] (A,
LAE). NEW CALEDONIA: Aoui, Mei, Baumann-Bodenheim 10156 (P), 10176 (A,
BRI, P); Bois du Mois de Mai, R. Blanche, Baumann-Bodenheim 14071 (A, BRI,P),
15179 (A, BRI, P); Col de Vulcan, Baumann-Bodenheim 8259 (A, BRI, P); Dambea,
Franc 3 (P), Franc 3A (L); Ermitage stream, Compton 149 (P); Mt. Paita, alt. 300 m,
Schlechter 14964, Oct 1902 (L, P, WRSL); Mt. Panie, Compton 1805, Aug 1914
(BM); Noudoure: MacKee 2670 (A, P); Yaouhe, Schlechter 14786 (P). SINE LOC:
Forster s.n. (BM); Vieillard 1227 (P); Schlechter 14787 (L, WRSL).
4. Piper bosnicanum C.DC. in Gibbs, Phyt. Fl. New Guinea, (1917) 207; Candollea
1 (1923) 221, as ‘bosniakanum’. Type: New Guinea, Schouten Islands, Bosnic, L.S.
Gibbs 6277 (holotype K; isotype L).
Glabrous climber. Lamina ovate, to 15 x 10 cm, with 3 pairs of lateral veins arising
directly from the base. Petiole to 2 cm long. Inflorescence to 5 cm long, shorter than
leaves; peduncular stalks c. 2 cm long, bracts oblong with sinuous margin. Female
flowers subsessile, spirally arranged; stigmas 3-4-fid, sessile, strongly reflexed. Fruits
subsessile, obovoid to ovoid, c. 10 x 6 mm broad, crowded.
Distribution: New Guinea and the Solomon Islands.
Notes: Piper bosnicanumiis very close to P. buruanum Migq. of Maluku: further study
is needed to assess their relationship.
Specimens examined: IRIAN JAYA: Schouten Islands, Bisnik, L.S. Gibbs 6277, Jan
(K; L); Vogelkop Peninsula, Aifact River, van Royen & Sleumer 7260, Nov (L). PAPUA
NEW GUINEA: Sepik Distr., Aitepe Subdistr., along Pieni River, Darbyshire &
Hoogland 8007, Jun (CANB). SOLOMON ISLANDS: Bougainville Island, north of
Buin, Craven & Schodde 256, Aug (CANB, LAE); Guadalcanal, Wanderer Bay, BS/P
9104 (LAE); Kolombangara, Bambari area, BSJP 751] (LAE); Malaita, Takwa, BS/P
10785 (LAE); New Georgia Group, Roniane Lagoon, BS/P 2918 (LAE); Wagina
Island, BSIP 5493 (LAE).
5. Piper lessertianum C.DC., J. Bot. 4 (1866) 164; Quisumbing, Philip. Jour. Sci.
18 Gard. Bull. Singapore 55 (2003)
43.1 (1930) 36, fig. 10, including var. oblongibaccum (C.DC.) Quisumbing.
Type: Philippines, Luzon, sine loc. Cuming 1343 (‘1342’), 1841 (holotype BM).
Synonyms: Chavica lessertiana Miq., Syst. Piperac., (1843) 270 non Piper lessertianum
C.DC. (1866): Piper pseudo-chavica C.DC. in DC., Prodr. 16.1 (1869) 351, nom.
superfl. Type: Philippines: Luzon, Cagayan Province, Cuming 1343 (holotype G, not
seen; isotype K).
Piper biroi K.Schum. & Lauterb., Nachtr. Fl. Schutzgeb., (1905) 238, syn. nov. Type:
New Guinea, near Malanaku, Biro 32, Oct. (holotype B).
Piper oblongibaccum C.DC. in Leafl. Philip. Bot. 3 (1910) 777, Type: Philippines:
Oriental Negros Prov., E/mer 9456 (not seen).
Piper lineatipilum C.DC., Nova Guinea Bot., 8.6 (1914) 1009, syn. nov. Type: New
Guinea, Mt. Hellwig, Rémer 962 (holotype L).
Piper viridibaccum Trel., J. Arn. Arb., 9 (1928) 150, syn. nov. Type: New Guinea,
Mowabula, L./J. Brass 1370 (holotype A, isotype BRI).
Climber, dioecious, glabrous to villose. Leaves shortly petiolate, petiole to 1 cm long.
Lamina narrowly ovate, ovate to broadly cordate, 12—32 x 5—17 cm; base unequally
rounded, one lobe acute to narrowly rounded, the other auriculate, lateral veins 34
pairs, the distal pair arising about 2 cm from the midrib, the others almost directly
from the base; apex often very long acute to acuminate. /nflorescences 5—12 cm long,
less than 5 mm thick; peduncle slender, 3—8 cm long, much longer than the petiole
and the floriferous part of the inflorescences. Male flowers with 2 subsessile stamens,
with pedicellate peltate bracts. Female flowers with 3-fid rounded sessile stigmas.
Fruits oblong, to 2.5 x 1.5 mm, borne crowded but not concrescent at maturity.
Distribution: Philippines, Sulawesi, New Guinea (Irian Jaya and Papua New Guinea).
Notes: This species is attributable solely to C.DC. who published it in 1866 basing on
Cuming 1342 (in herb. BM) without reference to Miquel’s earlier species Chavica
lessertiana which is based on Cuming 1343 (in herb. G). I am grateful to have the
advice of J.F. Veldkamp of Leiden (pers. comm., 2003) that Cuming 1342 in C.DC.’s
protologue of 1866 should read Cuming 1343 because Cuming 1342 refers to a species
of Gramineae. Nevertheless, the authority of Piper lessertianum (1866) remains
attributable solely C.DC. without reference to Miquel.
Later, in DC. Prodr. 16.1 (1869) 351, when C.DC. combined Miquel’s Chavica
lessertiana (1843) with his own Piper lessertianum (1866), he created the name Piper
pseudo-chavica for the combined species. By contemporary rules, the name Piper
pseudo-chavica is superfluous because his own Piper lessertianum (1866) is perfectly
legitimate and should have been used instead.
Malesian Piperaceae 3 19
Our Malesian species is not to be confused with Piper lessertianum (Miq.)
C.DC. in DC. Prodr. 16(1): 258 (1869) based on Ottonia lessertiana Migq., which
refers to an American species and, which binomial is a later homonym of our species.
Piper lessertianum C.DC. (1866) is readily recognised by the petioles being
very short and the peduncles very long, often more than five times longer than the
petiole and the floriferous part of the inflorescence. Yet, it is most variable in leaf
form with lamina ranging from very narrowly ovate with a long attenuate apex through
ovate ob-pentagonal to broadly cordate or heart-shaped. Besides, leaves borne on the
climbing parts are usually small and broadly cordate to heart-shaped while those
borne on the freely hanging branches tend to be ob-pentagonal to very narrowly ovate
with long attenuate apices.
All these leaf forms have now been observed to occur on the same plant as
evidenced by Vink & Schram BW 8&7]. In the circumstances, I have to assign to
synonymy Piper biroi, Piper lineatipilum and Piper viridibaccum as they are
distinguished purely on these leaf forms. For the same reason and after examination
of collections authenticated by Quisumbing, I am unable to maintain var.
oblongibaccum (C.DC.) Quisumbing.
Specimens examined: PHILIPPINES: Luzon, Cagayan Prov., Cuming 1343 (BM, K);
Mindoro, Mt. Halcon, Ramos & Edano 40676, Mar (NSW); Panay, Capiz Prov., Mt.
Bulilao, Martelino & Edano 35706, Jun (NSW); Jaminden, Ramos & Edano 30861,
31272, May-Apr (NSW); Libacao, Martelino & Edano 35395 (NSW), 35442, May-
Jun (NSW); Mt. Macosolon, Ramos & Edano 30776, Apr-May (NSW). SULAWESI:
Kabaena Island, Elbert 3393 (BO). IRIAN JAYA. Wati, Mapon Distr., /jiri & Niimura
674, Apr (L); Wissel Lakes, Doglia, Vink & Schram BW 8&71, May (CANB, LAE).
PAPUA NEW GUINEA: Milne Bay, near Mayu Island, Streimann NGF 28829, Apr
(L, NSW): Morobe Distr., Asubazo, Kairo NGF 24403, Jul (L); Northern Division,
Sibium Range, R. Pullen 5924, Sept (CANB).
6. Piper longipilum C.DC., Bot. Jahrb. 55: 216 (1918); Candollea 1: 206 (1923).
Type: New Guinea, Sepik, alt. 1-200 m, Ledermann 7565 (holotype B, isotype B).
Climber, dioecious, lightly pilose. Leaves shortly petiolate, petiole to 2.5 cm long.
Lamina ovate, c. 28 x 18 cm; base unequally cordate, lateral veins 5—6 pairs, the basal
pair arising c. 1 cm from the base; apex acuminate. Male inflorescences (immature) c.
8 cm long, c. 3 mm thick; peduncle slender, c. 1.7 cm long. Male flowers with
pedicellate peltate glabrous bracts. Female unknown.
Distribution: New Guinea.
Notes: The type material of this species consists of two sheets: the one with field
20 Gard. Bull. Singapore 55 (2003)
notes and annotation by C.DC in 1917 is the holotype, the other is the isotype. A
cursory examination of the type material suggests an alliance with Piper decumanum
L. of which it might represent a hairy variant. As the type seems to be the only
material there is of this species, I am unable to determine its status until I see more
collections, especially of female plants.
7. Piper melula Trel., J. Arn. Arb. 9 (1928) 148. Type: Papua New Guinea, Vaitata
River, L.J. Brass 1130 (holotype A; isotype BRI).
Dioecious glabrous climbers. Leaves moderately petiolate, petioles 2-4 cm long,
much longer than peduncles. Lamina irregularly ovate, broadly ovate to pentagonal,
c. 12 x 8 cm; apex bluntly shortly acuminate; base rounded to shallowly cordate;
lateral veins 3 pairs, the distal pair arising alternately from the midrib about 0.5—1 cm
from the base, the other 2 pairs directly from the base; only the distal pair reaching to
the apex, the 2" pair reaching to 1/2 the length of the lamina, the basal pair only to 1/3
of the length of the lamina. /nflorescences shorter than the leaves, males slightly
longer than females. Male floriferous part to 6 cm long on a peduncle to 7 mm long;
female floriferous part to 4 cm long on a peduncle to | cm long. Bracts circular,
margin sinuate, peltate, very shortly pedicellate. Male flowers with strongly reniform
anthers, dehiscing by long apical slits. Female flowers sessile, crowded; stigmas 3- to
4-fid, sessile, stigmatous part reflexed. Fruits almost entirely concrescent, free only
at the apices.
Distribution: Papua New Guinea.
Notes: Piper melula has the same leaf venation pattern as P. plagiophyllum K.Sch &
Laut., and in some cases even the lamina shape of that species, but it differs in being
a climber amongst other characteristics.
The infructescences of Piper melula appear similar to those of P.
pachyarthrum K.Sch., but these species differ in other respects.
Piper melula appears also to be related to P. elbertii C.DC. of the Lesser
Sunda Islands and P. austro-caledonicum of the Solomon Islands and New
Caledonia.
There is a collection, Kellman ANU 1597 (CANB) from Mindanao in the
Philippines, which is remarkably similar to Piper melula. However, as this collection
has a very young inflorescence, I hesitate to identify it as this species.
Specimens examined: PAPUA: Gulf distr., Tauri River, Schodde & Craven 4700,
Mar (CANB); Vailala River, L.J. Brass 1130, Mar (A, BRI); Milne Bay distr., Kwagira
River, Peria Creek, L.J. Brass 23997 (A, CANB, LAE), 24006, Aug (A, CANB, LAE).
Malesian Piperaceae 3 a |
8. Piper novo-guineense Warb., Bot. Jahrb., 13 (1891) 284, K.Schumann &
Lauterbach, Fl. Schutzgeb., (1900) 258. Type: New Guinea, Sattelberg bei
Finnschlafen, Warburg 20740 (holotype B; isotype A).
Glabrous climber. Lamina narrowly elliptic, narrowly ovate to broadly ovate or broadly
pentagonal, 17—19 x 5—9 cm; apex shortly blunt acuminate to long acute; base cuneate
to rounded, very slightly asymmetrical, not auriculate; lateral veins 3 pairs, the distal
pair arising slightly alternately from the midrib c. 1—2 cm from the base, the other 2
pairs arising directly from the base, sometimes another faint short pair arising from
the base; the veins on the underside, especially at the basal portion, often covered
with minute reddish brown scales, petiole to 1 cm long. Male inflorescences thin, as
long as the females, peduncle as long as the males. Male flowers with bracts as in the
females; stamens subsessile with broadly crescent-shaped anthers. Female
inflorescences to as long as the leaves, sometimes longer, usually to 15 cm long;
peduncular stalk c. 2 cm long. Female flowers sessile, stigma 3-4-fid. Fruits sessile,
oblong, crowded; bracts rounded peltate.
Distribution: New Guinea.
Notes: Piper novo-guineense is reported to be common in primary forest and older
secondary forests, between 500 and 2000 m altitude. At higher altitudes, the plants
tend to have smaller leaves.
This species is closely related to Piper macropiper Pennant and especially to
P. truncatibaccum C.DC.
Specimens examined: IRIAN JAYA: Lake Habbema, L.J. Brass 10299, Oct (A).
PAPUA NEW GUINEA: Morobe Distr., Boana Mt. Clemens 8467 (A); Gurokar, L.J/.
Brass 29499, May (CANB, LAE); Huon Peninsula, Mt. Rawlingson, Hoogland 9054,
Jun (A, BRI, LAE); Sattelberg, Warburg 20740 (B, A); Wantoat area, Kikiepa,
Womersley & Thorne NGF 11875, Jun (BRI, LAE); Selileo, Hellwig 580, Apr (B);
Sepik, near Kilifas village, Forman & Kumul NGF 48261, Mar (LAE).
9. Piper pallidilimbum C.DC., Nova Guinea Bot. 8.6 (1914) 1009. Type: New Guinea,
Mt. Hellwig, alt. 2600 m, von Romer 1316 (holotype L).
Synonym: Piper peracutilimbum C.DC., Nova Guinea Bot. 8.6 (1914) 1005, syn.nov.
Type: New Guinea, Mt. Hellwig alt. 2600 m, von Rémer 1252 (holotype L).
Climber, glabrous. Leaves shortly petiolate. Lamina ovate, 6-8 x 24 cm; base rounded,
apex acuminate. Lateral veins 2-3 pairs, distal pair arising from the midrib a little
22 Gard. Bull. Singapore 55 (2003)
above the base. /nflorescence shorter than leaves. Peduncular stalk 7-10 mm long,
about as long as petiole. Males up to 4 cm long, females c. 2 cm long. Flowers borne
crowded, sessile. /nfructescence congested, to 12 mm diam. as in Piper abbreviatum
Opiz.
Distribution: New Guinea.
Notes: The type material of Piper peracutilimbum is indistinguishable from specimens
of P. pallidilimbum. Therefore I see no reason to keep the two species apart.
Piper pallidilimbum is very close to P. abbreviatum Opiz from which it differs
in the lamina being furnished with fewer lateral veins, one of which issues from the
midrib, and also by the peduncular stalk being somewhat filamentous.
Specimens examined: IRIAN JAYA: Mt. Hellwig, alt. 2600 m, von Romer 1316 (L),
1252 (L); sine loc., Pulle 700 (L), 737 (L), 834 (L).
10. Piper pseudamboinense C.DC., Bot. Jahrb. 55 (1918) 206. Type: New Guinea,
Keneyia, alt. 150 m, Schlechter 15445 (holotype B).
Climber. Leaves very shortly petiolate. Lamina pentagonal to oblong, commonly c.
24 x 11 cm; apex bluntly acuminate, acumen to 1.5 cm long, base asymmetrically
cordate, one side often forming a large lobe, the lobes often covering the node; lateral
veins usually 5 pairs, the distal pair alternate, arising from the midrib c. 2—3 cm from
the base, the 2° pair often arising also from the midrib but very close to the base; the
other 3 pairs, often very short, issuing directly from the base; intercostals very
prominent; upperside of lamina glabrous, undersides with short brown pubescence
on the veins. Female inflorescences to as long as leaves, 20—24 cm long; the floriferous
portion as long or to only slightly longer than the peduncular stalk; peduncle glabrous.
Bracts circular, peltate, long pedicellate. Female flowers sessile, congested. Stigma
2-lipped, spreading, subsessile. Fruits sessile, free.
Distribution: New Guinea.
Notes: A lowland climber, Piper pseudamboinense 1s a little known species. Of the
two collections identified as this species, NGF 25678 appears to be the best match to
the holotype, the other being a more hairy variant.
Piper pseudamboinense is very closely similar to P. amboinense (Miq.) C.DC.
and from which it differs in the more prominent lateral veins and its stigma being
2-lipped. It is also closely similar to P. lessertianum C.DC. from which it can be
distinguished by the 2-lipped stigma in addition to lamina and inflorescence characters.
Malesian Piperaceae 3 23
Specimens examined: PAPUA NEW GUINEA: Morobe District, Buso River, NGF
25678 Aug (CANB, LAE); Oomsis Creek, T.G. Hartley TGH 10564, July (A, CANB,
LAE); Keneyia, alt 150 m, Schlechter 18445 (B).
11. Piper reinwardtianum (Miq.) C.DC., DC. Prodr. 16.1 (1869) 354. Merrill, Inter.
Rumph. Herb. Amb., (1917) 182. Basionym. Macropiper reinwardtianum Miq..,
Linnaea 21 (1848) 481. Type: Indonesia, Maluku, Nussa Laut, Reinwardt, July 1821
(holotype L?).
Synonym: Chavica reinwardtiana (Migq.) Miq., Ann. Mus. Bot. Ludg. Bat.1
(1863) 136.
Dioecious shrub, glabrous to lightly pilose. Leaves with petiole 2.5—5 cm long. Lamina
broad ovate to rounded, c. 16 x 15 cm, slightly asymmetrical; apex shortly acute, base
asymmetrically truncate-cordate, lateral veins usually 34 pairs, upperside of lamina
glabrous, undersides lightly pilose on the veins. /nfructescences to 11 cm long, peduncle
glabrous to 2.5 cm long. Female flowers sessile, congested. Stigma 3, rarely 4, short
thickset. Fruits sessile, somewhat obovoid, concrescent on the lower half.
Distribution: Indonesia, Maluku.
Notes: I have not seen any type material of this species which is probably in the
Leiden Herbarium to which Reinwardt donated his collections from the region. The
collections of Robinson and Teysmann cited herewith are all that I have seen of this
species; and I must of neccesity rely on C. de Candolle and Merrill in the interpretation
of this apparently rare species.
In general growth habit, Piper reinwardtianum recalls that of P. sundaicum
Blume, also a shrub from the same region. However, it has infructescences similar to
P. mestonii Bailey of New Guinea and Australia from which it can be distinguished
by the stigma being 3- or 4-fid, not bifid, and the plant a shrub, not a climber. In these
respects, P. reinwardtianum appears to be an interesting species, and further
investigation may yield insight into the phytogeograpical relationship between the
three species.
Specimens examined: INDONESIA: Maluku, Ambon, Robinson 60 (L). Ceram,
Teysmann s.n. (L).
12. Piper subvirosum C.DC., Bot. Jahrb. 55 (1918) 215. Type: New Guinea,
Alexishafen, Wiesenthal 68 (holotype B).
24 Gard. Bull. Singapore 55 (2003)
Climber, twigs minutely pubescent. Lamina broadly ovate, 15—20 x 10-12 cm, upper
surface glabrous, lower surface sparsely covered with short stiff hairs especially on
lateral veins on basal part of the lamina; apex shortly acuminate, acumen 1.5—2 cm
long; base broadly cuneate, rounded to truncate; lateral veins 3 pairs, distal pair
alternate, arising from midrib a little above the base, the others directly from the
base; intercostals prominent. Petiole to 2 cm long. Female inflorescence to 27 cm
long, peduncle to 3 cm long, floriferous part to 24 cm long; inflorescences usually
longer than leaves, peduncle longer than petiole; mature inflorescence to 7 mm diam.
Female flowers sessile, not concrescent, with 3- or 4-fid stigmas.
Distribution: New Guinea.
Notes: Piper subvirosum is related to P. macropiper Pennant from which it can be
distinguished by the much larger leaves and inflorescences, at least one pair of the
lateral veins arising from the midrib, the lamina base not auriculate, and the peduncle
not excessively longer than the petiole.
Not much is known about this species. The two recent collections cited here
are the closest match to the holotype.
Specimens examined: IRIAN JAYA: Star Mts., Sibil Valley, alt. 1200-1500 m,
Kalkman & Nicholas 4160, May (L). PAPUA NEW GUINEA: Central Division, Mt.
Tafa, L.J. Brass 4127 (A); Alexishafen, Wiesenthal 68 (B).
13. Piper sundaicum Blume, Verh. Bat. Genoots. 11 (1826) 187, fig. 14; C.DC.,
DC. Prodr. 16.1 (1869) 352; Koorders, Verh. Kon. Ak. Wet. Amsterdams. 2, 14.4
(1908) 42, reprint; Koorders, Exkursionsflora van Java. 2 (1912) 26; Koorders-
Schumacher, Exkursionsflora Atlas. 4.2 (1924) 445, fig. 718G. Type: Maluku, Ambon.
Blume s.n., Leiden Herb. 908.363-43 (lectotype L, here designated).
Synonyms: Chavica sundaica (Blume) Mig. Syst. Piperac., (1843) 274; Fl.Ind. Bat
1(2) (1859) 446, exclusive of the synonym P. a/bum Vahl.
Piper amplilimbum C.DC., Candollea 1 (1923) 213; Schroeder, Candollea 3 (1926)
135; Chew, Blumea. 20 (1972) 145, fig. 1, syn nov. Type: Maluku, Ambon, Teysmann
s.n. 1905 (holotype BO).
Dioecious scandent shrubs, entirely glabrous. Twigs 5—8 mm diam.; internodes smooth;
stipular scars prominent. Lamina symmetrically ovate, (23—)25—27 x 14-19 cm,
somewhat coriaceous; apex acuminate, acumen to 2 cm long; base rounded; margin
entire; lateral veins 34 pairs, very prominent, the distal pair arising alternately from
the midrib at 2-3 cm from the base, the others directly from the base; intercostals
Malesian Piperaceae 3 25
numerous, not straight, quite prominent. Petioles c. 2 cm long, smooth. Stipules to 8
x c. 1 cm, apparently not adnate to petioles. /nflorescences: male long, thin; female
11-13 x 0.7 cm, the peduncular stalk c. 1.5 cm long, glabrous. Bracts somewhat
irregularly circular, peltate. Female flowers sessile; ovary pyramidal; stigmas 3-fid,
short, sessile, slightly reflexed. Fruits ovoid, sessile, c. 3 cm long: crowded, not
concrescent.
Distribution: Indonesia, Maluku.
Notes: Blume recorded this species as also occurring in Java. Koorders (1908, 1912)
doubted the Javanese record as he failed to find any collection from that island. I have
also been unsuccessful in finding any material of the species from Java. This is probably
a very rare species: the type materials are all that is known of this species.
I have compared the type materials of Piper sundaicum with that of P.
amplilimbum C.DC., and I am unable to distinguish the one from the other.
Piper sundaicum appears to have some relationship with Piper reinwardtianum
from the same region (see above).
Piper sundaicum has been identified with P. album Vahl by previous botanists.
The two species are however, quite distinct: P. album Vahl is a climber, our species is
not.
Specimens examined: MALUKU: Ambon, Blume s.n. (L): Teysmann s.n. 1905 (BO).
Acknowledgements
I thank the curators of the following herbaria (identified by their acronyms) for loan
of their collections, including type materials, for my study at the Herbarium of the
Royal Botanic Gardens, Sydney where I held tenure as Senior Research Scientist
from 1970 to 1975: A, B, BO, BRI, CANB, K, L, LAE, P, PR, PRC and WRSL.
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Garden’s Bulletin Singapore 55 (2003) 27-30
A New Species of Schismatoglottis (Araceae) from
Sabah, Malaysia
A. HAY AND C. HERSCOVITCH
Royal Botanic Gardens, Mrs Macquarie’s Road, Sydney, NSW 2000, Australia
Abstract
Schismatoglottis tahubangensis A. Hay & Herscovitch is described, illustrated and inserted
into the latest key to Bornean Schismatoglottis species.
Schismatoglottis tahubangensis A. Hay & Herscovitch, sp. nov.
A Schismatoglottis platystigma M. Hotta spadice tenuiore, staminodiis inter pistilla,
connectivo longiore differt. - Typus: Malaysia, Sabah, just outside Kinabalu Park,
100 m to Tahubang River along route to Marai-Parai, 11,000 m alt. [sic], 12 Sep
1993, J. Nais, Dolois, Harry & Masius SP 05312 (KEP, holo).
Terrestrial herb c. 30 cm high. Stem creeping or decumbent, rooting between the
nodes, c. 1 cm diam., with internodes c. 5 mm long. Leaves 6 together; petiole 10-17 cm,
sheathing in the lower 1—2 cm, wings of sheath extending into a papery narrowly
triangular ligule to 2.5 cm; blade 11—15 cm long, 3.5—5 cm wide, narrowly obovate,
somewhat asymmetric, base truncate to acute, somewhat asymmetric, inserted into
adaxial face of petiole, apex acute; midrib pronounced and minutely puberulent
abaxially, with c. 10 primary lateral veins on each side diverging at c. 45°, interprimary
veins somewhat conspicuous abaxially, obscure adaxially, fine. /nflorescence solitary,
subtended by an inconspicuous papery cataphyll c. 6 mm long; peduncle 3 cm long.
Spathe pinkish, 7 cm long, slender, weakly constricted; lower spathe 1.5 cm long,
3mm diam. Spadix very slender, cylindric for most of its length, 5.3 cm long, 1.5 mm
diam.; female zone 1.2 cm long, (not clear whether adnate to spathe at base); pistils
closely packed, subglobose, 0.3 mm diameter; stigma discoid, almost as wide as the
ovary; interpistillar staminodes conspicuous, round and flat-topped or concave, drying
pale yellow, of the same height and diameter as pistils; sterile interstice c. 2 mm long
comprised proximally of 2 irregular whorls of looser packed, 0.5 mm diam., flat-topped
staminodes, differentiated from interpistillar staminodes by larger size, slightly convex
top, and dark yellow colour on drying, and distally of 2—3 irregular whorls of sterile
28 Gard. Bull. Singapore 55 (2003)
stamens drying dark brown; male zone 1.2 cm long; stamens subsessile, more or less
oblong ovate from above, more or less irregularly paired and oriented along long
axis of spadix, c. 0.8 mm wide; thecae round, opening by a single apical pore;
connective robust, columnar, flat-topped and somewhat impressed, slightly exceeding
the thecae in height and about 1.5 times their diameter, rounded triangular from
above with ends indented above thecae; appendix 2.8 cm long, tapering to a fine
naked tip, composed of flat-topped, more or less rounded or irregularly polygonal
staminodes c. 0.3 mm diam. similar to those of sterile interstice. Fruit unknown.
Distribution: Malesia — Borneo, Sabah, Kinabalu (known only from the type).
Habitat: Ona steep slope in unspecified vegetation type at c. 1,100 m alt. [Note that
the label on the type records the altitude as 11,000 m, which 1s, of course, impossible].
Notes: This species somewhat resembles Schismatoglottis platystigma (Brunei) in
having discoid stigmas and relatively small thecae on the edge of a broad connective.
However, it is readily distinguished from that species by the presence of interpistillar
staminodes, the subsessile stamens with a much higher connective and the more
slender spadix.
Schismatoglottis tahubangensis may be inserted into the key to Bornean
Schismatoglottis provided by Hay & Yuzammi (2000) as follows:
17a. Inflorescence solitary or two together; anthers with broadly dilated connective
Fe ee EO RES re ree ree Teeter en ee 18
17b. Inflorescences c. 4-6 together; anthers dumb-bell-shaped, with the connective
a i a a ee a ERE NITED
18a. Interpistillar staminodes absent; appendix 1—2 cm long, blunt-tipped; Temburong,
BOE? 2A ANE. RU TER G.. RUA ASS ORAS Se
18b. Interpistillar staminodes numerous; appendix c. 3 cm long, tapering to a fine
CIN, PA MNANAIEL, DOIN. os os nog om eee i rmadtaesapasenn esata as <email S. tahubangensis
19 etc = 18 etc in Hay & Yuzammi (2000).
Schismatoglottis tahubangensis 29
Figure 1. Schismatoglottis tahubangensis Herscovitch & A. Hay
A. habit; B. ligule at base of petiole; C. detail of leaf venation; D. midrib detail; E. inflorescence with
part of spathe removed: F. pistils with interpistillar staminodes; G. sterile interstice with staminodes and
sterile stamens; H. stamens; I. staminodes of appendix. J. Nais, Dolois, Harry & Masius SP 05312.
30 Gard. Bull. Singapore 55 (2003)
Acknowledgements
We thank Josef Bogner for drawing our attention to the specimen on which this species
is based, Dan Nicolson for commenting on the ms and and Lesley Elkan for the
illustration.
Reference
Hay, A. & Yuzammi. 2000. Schismatoglottideae (Araceae) in Malesia I —
Schismatoglottis. Telopea. 9: 1-177.
Garden’s Bulletin Singapore 55 (2003) 31-34
An Unusual New Species of Homalomena (Araceae) from
New Guinea
C. HERSCOVITCH AND A. HAY
Royal Botanic Gardens, Mrs Macquarie’s Road, Sydney, NSW 2000, Australia
Abstract
Homalomena impudica Herscovitch & A. Hay is described, illustrated and inserted into the
latest key to Homalomena species of New Guinea.
Homalomena Schott was revised for New Guinea by Hay (1999). An overlooked
specimen on loan to NSW from LAE represents a very distinctive undescribed species.
Homalomena impudica Herscovitch & A. Hay sp. nov.
Homalomena tenuispadix Engl. simulans sed inflorescentia femina spatham aequanti,
inflorescentia mascula clavata tota exserta differt. - Typus: Papua New Guinea,
West Sepik Province, Vanimo, Ossima, 28 Jan 1969, H. Streimann & A. Kairo NGF
39239 (holo, LAE).
Erect multi-stemmed herb to c. 30 cm high. Stem condensed, 1.5 cm diam. Leaves to
8 together; petiole longer than blade, 17—32 cm long, glabrous, slender, sheathing in
lower 6—7 cm; blade dull green above, green below, glabrous, sagittate to ovate-
cordate,14-19 cm long, anterior lobe 12—16.5 cm, c. 8 cm wide, posterior lobes
triangular to rounded, 2.5—4 cm long; midrib more prominent abaxially, with c. 8
pairs of lateral veins diverging at c. 45 °. Inflorescences to 9 together, each subtended
by a narrow triangular papery cataphyll to 4 cm long; peduncle erect, later declinate,
4.5—6 cm long, longer than spadix. Spathe slightly constricted in bud, then narrow
cylindrical, 2.0—2.8 cm long, 5 mm diam., with a conspicuous rigid darker sometimes
reflexed apiculum 2—3 mm long. Spadix 2.7—3.6 cm long, much exceeding the spathe
at anthesis, stipitate for c. 5 mm; female zone equalling to exceeding the spathe, 2.0—2.4
cm long; pistils very widely scattered along axis of female zone, becoming loosely
and irregularly whorled distally, globose, c. 1 mm diam., infrapistillar staminodes
absent; stigma button-like, sessile, 0.3 mm across; male zone narrow oblong-ovoid, 0.7—1.1 cm
32 Gard. Bull. Singapore 55 (2003)
long, whitish, fully exserted from spathe; male flowers (-)2-merous; stamens more-
or-less oblong to irregularly polygonal from above, c. 1 x 0.3-0.5 mm, the thecae
overtopped by connective. Fruit unknown.
Distribution: Malesia— New Guinea, Papua New Guinea, West Sepik Province (known
only from the type locality).
Habitat: In a dry stream bed in lowland rain forest at c. 30 m alt.
Notes: This species resembles Homalomena tenuispadix Engl., but differs in the greatly
elongated and lax female zone which equals to exceeds the spathe, and in the
completely exserted and somewhat clavate male zone. The inflorescence at first
resembles that of Schismatoglottis after the male zone has been exposed by the fall of
the spathe limb, until it is noted that the spathe, as typical for Homalomena, 1s fully
persistent.
While some New Guinea species of Homalomena, such as H. davidiana A.
Hay and others (see Hay 1999), characteristically have the male zone partially exserted ~
from the spathe after anthesis, this species extends the entire male zone beyond the
apex of the spathe prior to anthesis. This characteristic, as distinct from the state
where the male zone is exposed by the spathe being wide open ms anthesis, appears
not to be recorded for any other species in the genus.
Field notes describe the flower (sic) as white and the fruit green. The specimen
includes fragments of declinate fruiting peduncle, but the infructescence is not present.
It is unclear whether or not the colour observations refer to the spathe at different
stages.
The specific epithet, meaning lewd or shameless, alludes to the fully exposed
male zone of the spadix.
Homalomena impudica can be inserted into the key provided by Hay (1999) as follows:
10a. Spathe tip + hood-forming; spadix contained within the spathe after anthesis;
peduncle tice Sorter tian tite PetiOles .2-. tcc cce cee ree ececee se H. lauterbachii
10b. Spathe with the tip reflexed; spadix exserted from the spadix during and/or after
anthesis; peduncle c, 1/3—1/2 the leneth of (he petioles |. na beriecekes Gannann een 11
lla. Spadix with the male zone clavate and completely exserted from the spathe at
anthesis). A SS a ee eae See H. impudica
1 1b. Spadix with the male zone cylindric-tapering, not exceeding the spathe at anthesis,
partially exserted from spathe after anthesis ...................ssseeeseeeH. Cenuispadix
12 etc. = 11 etc. in Hay (1999)
Homalomena impudica 33
Figure 1. Homalomena impudica Herscovitch & A. Hay.
A. habit; B. inflorescence with part of spathe removed; C. tip of spathe; D. inflorescence in bud; E.
pistils; F. stamens. Streimann & Kairo NGF 39239.
34 Gard. Bull. Singapore 55 (2003)
Acknowledgements
We thank LAE for the loan of the specimen on which this species is based and Catherine
Wardrop for the illustration.
Reference
Hay, A. 1999. Revision of Homalomena (Araceae-Homalomeneae) in New Guinea,
the Bismarck Archipelago and Solomon Islands. Blumea. 44: 41-71.
Garden’s Bulletin Singapore 55 (2003) 35-60
Pleuroschisma, a New Section of Cyrtandra (Gesneriaceae)
from Borneo
O.M. HILLIARD’, B.L. BURTT! AND M.H. BOKHARI?
‘Royal Botanic Garden Edinburgh, EH3 5LR, U.K.
*Multan, Pakistan
Abstract
Twelve Bornean species of Cyrtandra constitute a new section, Section Pleuroschisma, the
name derived from an important diagnostic character, namely the unique fruit with a median
septicidal split on each side (not reaching either apex or base of the fruit). Nine of the species
and two varieties are newly described, C. angustielliptica Hilliard & B.L.Burtt, C. coacta
Hilliard & B.L.Burtt, C. insolita Hilliard & B.L.Burtt, C. linauana B.L.Burtt, C. seganica
Hilliard & B.L.Burtt, C. tesselata Hilliard & B.L.Burtt, C. tunohica Hilliard & B.L.Burtt, C.
pendulifera Kraenzl. var. grossipilosa Hilliard & B.L.Burtt and C. sarawakensis C.B.Clarke
var. Jongipilosa Hilliard & B.L.Burtt. Two species recognized remain nameless because the
material is inadequate to typify a name.
Introduction
There are already 40 sectional names in Cyrtandra and the limits and interrelationships
of the groups so designated are far from being understood; the addition of yet another
sectional name to the list may seem somewhat irresponsible. Nevertheless, the fruit
characters that are used to diagnose this section - the smooth fruit wall (due to a thin
layer of sclerenchyma beneath the epidermis, Fig. 3), and the septicidal splits with
inrolled margins (Fig. 1) form a unique combination in this small group. It is the
lateral splits in the fruit-wall that form the basis for the sectional name Pleuroschisma;
these splits, it should be noted, are septicidal and there is some reason to suppose that
septicidal dehiscence may have been the primitive condition in the family as it is
found in Ramonda, Haberlea, in the type species of Corallodiscus, all on the northern
fringe of the subfamily Cyrtandroideae, and in the tribe Coronantherae, a basal tribe
of subfamily Gesnerioideae found in New Caledonia, E. Australia, New Zealand and
southernmost S. America. Its presence elsewhere in the family (e.g. in Leptoboea)
may be secondary. It must be noted, however, that the lateral splits in this new section
of Cyrtandra do not reach from apex to base of the fruit (Fig. 1) and do not give rise
to free fruit-valves, as in a fully septicidal fruit. These features set section smaller.
36 Gard. Bull. Singapore 55 (2003)
Figure 1. Mature fruit of Cyrtandra sarawakensis
The two halves remain held together at apex and base. Burtt B4¢71]2 x4.
Pleuroschisma apart from all the other sections.
As so often in herbarium work on Cyrtandra, we have been much hampered
by lack of flowers on many specimens, although mentioned in the collector’s field
notes. Nevertheless, it is not the flowers that provide the diagnostic features of the
Section: it is the fruits. We have therefore felt justified in naming C. tesselata (no. 6),
C. seganica (no. 7) and C. insolita (no. 11), which we know only in fruit. In the eight
species where we have seen flowers, the major floral characters are uniform: calyx
divided almost to the base into five deltoid lobes, corolla c. 35-40 mm long, white,
cream or palest yellow with two orange/yellow bars in the throat, puberulous outside,
disc cupular, occasionally deeply excavated on one side, ovary crowned with a coma
of relatively long hairs, otherwise so minutely gland-dotted or pustulate as to appear
glabrous on casual inspection, style puberulous, stigma distinctly bilobed, lobes
relatively large, clavate in outline.
Leaf-venation provides a useful character in distinguishing some of the species,
both the number of lateral veins on each side of the midrib, and the pattern described
by the tertiary venation, of which three main types have been distinguished; for these
we use the terms subscalariform, reticulate and pinnate (Fig. 2, A,B,C,D). In some
species, the tertiary venation is invisible (e.g. C. insolita).
Cyrtandra sect. Pleuroschisma af
Figure 2. Types of tertiary venation found in Cyrtandra sect. Pleuroschisma
A. subscalariform (C. sarawakensis, Burtt B1947, about actual size). B. finely reticulate (C. tesselata,
Hotta 1417, slightly enlarged). C. coarsely reticulate (C. sp. nov. no. 12, Burtt B2658, x2). D. pinnate
(C. coacta, Beaman 10656, slightly enlarged).
38 Gard. Bull. Singapore 55 (2003)
The distinctive leaf anatomy, with tracheoids in the hypodermis, has already
been illustrated (Bokhari & Burtt, 1970, Plate 2, G,H): the tracheoids may have either
parallel or reticulate bars of thickening (Fig. 4). In some species (e.g. C. hoseana),
similar tracheoids are found in the mesocarp of the fruit; spheroidal sclereids are
commonplace and abundant (Fig. 3), making fruit-sectioning very difficult. In several
species, polymorphic sclereids occur in the mesophyll of the leaf (C. tesselata, C.
seganica, C. hoseana, C. angustielliptica); in C. penduliflora, they may be present or
absent. The other species in section Pleuroschisma lack sclereids. (C. insolita and sp.
no. 12 have not been examined).
Protection of the apical bud in Cyrtandra is achieved in several ways. In
section Pleuroschisma the bud is hidden between the erect petioles of the two fully
developed uppermost major leaves on the stem. The following observations were
made on herbarium material, and observations are needed on living plants to determine
time-intervals in leaf development: when the developing leaf in C. /inauana measured
38 mm, the young apical bud was only 2.5 mm long. We examined the stem apex of
C. linauana, removing the tip of the stem and rehydrating it. The very young major
leaf of the apical bud is hidden by the conduplicate base of the minor (stipule-like)
leaf of the uppermost pair of fully developed leaves. It is plane, measured 38 x 8 mm
including the petiole (scarcely developed at this stage), and both surfaces are thickly
clad in long silky hairs. The minor leaf has similar indumentum and 1s well developed,
being 20 mm long. In contrast, the fully developed leaves below the apex show no
trace of the juvenile indumentum except over the midrib and lateral veins on the
lower surface. There is the same abrupt transition from a very small juvenile leaf to
a fully developed one in five other species where we had stem-apices suitable for
inspection (C. penduliflora, C. sarawakensis, C. coacta, C. angustielliptica, C.
insolita). These species in section Pleuroschisma are all strongly anisophyllous. A
comparable condition in isophyllous species was noted by Burtt (2001, p.399).
Field observations by one of us (B.L.B.) found that in C. hoseana and C.
penduliflora, as the new apical leaf begins to grow out, the petiole is held erect but the
young lamina is pendulous, the hairs in C. hoseana being silvery white, while in C.
penduliflora they are dark red. As the leaf blades enlarge, the hairs become more
spread out, and are eventually shed, resulting in the blade of the mature leaf being
glabrous above. Whether or not these young leaves perform any attractive function is
not known, but they certainly catch the eye of the plant-hunter.
Cyrtandra section Pleuroschisma Hilliard & B.L.Burtt sect. nov. ab sectionibus
omnibus ad huc descriptis fructibus duris laevibus (strato annulari subepidermali
schlerenchymatis uni- vel bi-seriato), parietibus in medio longitudinis fissuris duabus
septicidalibus marginibus involutis notatis, foliis cellulis tracheoidalibus
hypodermalibus praeditis facile distinguenda.
Cyrtandra sect. Pleuroschisma
WN |
|
39
Ay
4 4
ie!
=
~ i
a> Ss
——
——— ~~ —
~
Figure 3. Transverse sections of part of fruit wall and septum
A. Cyrtandra sarawakensis; B. Cyrtandra penduliflora. b. scl. = brachysclereid; fib. 1. = fibrous layer;
m. scl. = macrosclereid; scl. p. = sclerified parenchyma [sclerenchyma]; sp. scl. = spheroidal sclereid; tr.
= tracheoid [with reticulate thickening]. M.H. Bokhari del.
Figure 4. Sclereid and tracheoid types.
A. polymorphic sclereids. B. tracheoid with reticulate bars. C. tracheoid with parallel bars. M.
Mendum after pencil sketches by M.H. Bokhari.
40 Gard. Bull. Singapore 55 (2003)
Species typica: C. sarawakensis C.B.Clarke
Distribution: Endemic to Borneo.
Simple-stemmed perennial herbs (C. insolita bushy), leaves usually strongly
anisophyllous, tracheoids, as far as is known, in hypodermis, inflorescence a dichasial
cyme, bracts free, bracteoles present or absent, calyx divided almost to base, corolla
medium-sized, white to palest yellow ground colour, disc cupular, ovary crowned
with a coma of relatively long hairs, style pubescent, stigmatic lobes large, clavate,
fruit with two septicidal grooves (later splits).
Key to Species
la. Lateral veins in largest mature leaves up to 10 each side of midrib ............... 2
ib, ‘Lateral veins 14-20 on each side of madrid... a eee ~
2a. Hairs (lower leaf surface) on midrib and lateral veins very inconspicuous,
short (to | mm), strongly appressed, pedicels c. 20 mm long,calyx lobes
10-15 tats... eae ee a ee ee 8. C. linauana
2b. Hairs (lower leaf surface) on midrib and lateral veins long, silky, strongly
appressed, sometimes matted together to produce a papery surface,
pedicels c. 5—8 mm, calyx lobes to'8 mii) ree eee 3
3a. Largest leaves c. 60-165 mm broad, petioles 75-135 mm, bracts c. 35— 40 x
10—16 mm, conspicuous, stronely veined. ::... 2.25.2... ae 3. C. coacta
3b. Largest leaves c. 36-60 mm broad, petioles 40-45 mm, bracts c. 5.5—7 x
1.8—2 mm, inconspicuous, only midrib visible.............. 10. C. angustielliptica
4a. Hairs (lower leaf surface) on midrib spreading: .... 200, clk. a. ..s0ek-seeeeesct mean 5
4b. Hairs (lower leaf surface) on midrib strongly appress................::sssecceeeeeeeees 10
5a. Peduncle c. 45-240 mm long, inflorescence lax, trailing..............eeeeeseeeeeeeeeeeeee
Pie: omy te 1. C. penduliflora var. grossipilosa
5b. Peduncle c. 2-30 mm long, inflorescence neither lax nor trailing.................. 6
6a. Bracts c. 20-50 x 7-25 mm, 3-5 veins, clearly visible, calyx lobes
2A MAREE TOM ees Sead St ae a a leek ae eee 7
6b. Bracts c.15—18 x 2—3 mm, only midrib visible, calyx lobes
©. 6-10 TM TOTS oie occ eccce deed, Din, babdnetdaetesncs eee ttt. Aa state ge en ee 9
7a. Inflorescence a several- to many-flowered cyme, both
bracts and bractéolés present... ...050..0. 00. 82. ee eee 8
7b Flowers 1-3, clustered and, at least when young, hidden by two
foliaceous bracts, bracteoles wanting. ...............cssssssseeeeeeeeeeeeeeees 11. C. insolita
8a. Petioles 80-150 mm long, peduncles 5—30 mim...............cccceecesceeeeeeeeeeeeeeeeeeeeeeeees
sauaind adboonvivesniNootecd seed dadana easel pl ee ee 2. C. sarawakensis var. longipilosa
Cyrtandra sect. Pleuroschisma 4]
8b. Petioles 30—70 mm long, peduncles 0—2 mm............... 5. Cyrtandra sp. nov.
9a. Largest leaves c. 26-34 mm broad, lower surface displaying finely
reticulate tertiary venation, petiole c. 20—25 mm long ............ 6. C. tesselata
9b Largest leaves c. 85—110 mm broad, lower surface displaying coarsely
reticulate venation, petiole c. 45—50 mm long........... 12. Cyrtandra sp. nov.
10a. Peduncles c. 50-175 mm long, inflorescence lax, trailing...
MPT eisintex. 0k). neo) er, SL. BAO... ae ee 1. C. penduliflora
10b. Peduncles c. 1-30 mm long, inflorescences neither lax nor
mbit os Be es, a EO) 3 be ade RL a ail WR 11
ibaai ise. 60-20 ann booed cece UA heck coe tO RR ES 12
RAG ) aeaeee ond 1S bread a3 41h. een ahs; ae id is en NII ‘3
12a. ‘Tertiary venation on lower leaf surface subscalariform, hairs on
stem, leaves (including petioles), bracts, bracteoles, pedicels and calyx
appressed, all: hairs. separatel ncveli icc egl. wisn Ab sdae des 2. C. sarawakensis
12b. Tertiary venation more or less pinnate, hairs on all parts (as above)
matted together giving a papery look to the surface................... 3. C. coacta
13a. Stem silky-villous, tertiary venation (lower leaf surface) coarsely
reticulate, almost invisible, largest mature leaves c. 85—95 mm broad,
foisbengthiz breath wo 0e3G: beniys. hd enh Qed 9. C. hoseana
13b. Stem pubescent, tertiary venation more or less subscalariform,
leaves c. 35-60 mm broad, ratio length: breadth 4—-5.7:1........ eee 14
14a. —_ Largest leaves c. 35-50 mm broad, lateral veins 13—15 on each side
of midrib, bracts 20—30 x 4—7.5 mm, at least midrib clearly
ie aa oe eet hires. Dada. heed Sao r9 tate. a RSI 4. C. tunohica
14b. — Largest leaves c. 55-60 mm broad, lateral veins 18 on each side
of midrib, bracts c. 20 x 4 mm, midrib scarcely
ae ry ee ie Behar tit osiencssisvedass oonssanccsievcassse 7.C. seganica
1. Cyrtandra penduliflora Kraenzl., Mitt. Inst. Allg. Bot. Hamburg 7:100
(1927); B.L. Burtt, Notes Roy. Bot. Gard. Edinb. 30:36 (1970).
Lectotype (Burtt, 1970): Kalimantan, Bukit Obat [c.0°56'N 113°20'E], 150m,
Winkler 1328 (lecto HBG, isolecto E).
Unbranched herb, stem to c. 600 mm long, 9-16 mm in diam., base often decumbent,
rooting, prop roots present, young parts strongly appressed-pubescent. Leaves
opposite, strongly anisophyllous, few forming a fan at apex of stem, reduced leaves
stipule-like, c. 20-46 x 5-14 mm, lanceolate, acuminate, strongly appressed-
pubescent, largest developed leaves c. 210-300 x 150—200 mm, very broadly elliptic,
apex abruptly acute, base cuneate, very narrowly decurrent, briefly or to c. 40 mm,
margins entire to obscurely to more distinctly irregularly serrate, lateral veins 1 1—13
on each side of midrib, tertiary venation more or less pinnate, upper surface glabrous
4? Gard. Bull. Singapore 55 (2003)
at maturity, coarsely pitted, lower surface with fine appressed hairs on veins and
blade; petiole 130-210 mm long, hairy as midrib. /nflorescence an axillary very lax
dichasial cyme, loosely branched, trailing, tending to spring from axils of fallen leaves,
peduncle c. 50-175 mm long, pubescent. Bracts (lowermost pair) c. 15—57 x 4-18 mm,
lanceolate, acuminate, finely pubescent. Pedicels 6-20 mm long, puberulous. Calyx
5-lobed almost to base, lobes subequal, c. 6-14 x 1.5 mm, deltoid, outside puberulous,
inside minutely gland-dotted. Corolla c. 37 mm long (no complete flowers seen,
only a big bud), white with an orange-yellow blotch or two bars in throat, puberulous
all round mouth. Stamens inserted c. 15 mm above base of tube, filaments c. 5 mm
long, glabrous, anthers 3.5 x 1.8 mm, cohering face to face by a small ligature,
connective glabrous or a very few glandular hairs. Disc 1 x 1.8 mm, cupular. Ovary
6 x 1 mm, minutely pustulate, coma of hairs at apex. Sty/e 12 mm, glandular-puberulous.
Stigmatic lobes c. 1.5 x 1 mm (will enlarge), clavate. Fruit c. 20-45 x 5~7 mm, pericarp
smooth. Seeds c.0.3 x 0.25 mm, testa red-brown.
Notes: Cyrtandra penduliflora is a plant of the forest floor, often on steep banks, the
stem supported by prop roots. It is distinctive by virtue of its trailing inflorescences,
the peduncles mostly very long, the cyme always very lax, far laxer than in any other
species of this group. The leaves are broad in relation to length, those of the type
specimen, collected in southern Kalimantan, measuring roughly 300 x 200 mm, and
exactly matched by those of Burtt 2610 from Sarawak collected 1° of latitude north
of the type collection and at very nearly the same degree of longitude (1°56’N 113°06’E
versus 0°56’N 113°20’E), but nearly all specimens have the leaves very broad in
relation to their length.
Tracheoids with parallel bars of thickening occur in the hypodermis;
polymorphic sclereids are often present in the spongy mesophyll but they may be
absent.
Other specimens examined: Sarawak. Miri distr., Niah Forest Reserve, [c. 3°52'N
113°44'E], Anderson $31663 (E). Bintulu distr., Ulu Sungai Sinonok, 20-60 m, Hotta
14247 (E, KYO). Tatau, Batang Anap, Ulu Sg. Kana [c. 2°54'N 112°50'E], Mokhtar
& Jugah 41779 (E). Sungai Kakus [c. 2°48'N 112°42'E], 30-80 m, Hirano & Hotta
220 (E, KYO). Ulu Kakus [c. 2°48'N 112°42'E], Anap, Haron S29980 (K, E). Ulu
Segan [c. 2°30'N 113°E], N Setungan, c. 50 m, Ashton $22019 (E). Pelagus Rapids
on Rajang [‘Rejang’], c. 2°10'N 113°E, Burtt & Woods B2559 (E). Sungai Bena [c.
1°56'N 113°06'E], a tributary of Sungai Sut, Burtt B26/0 (E). Sungai Jelok, near
Bukit Sengkajang [1°14'N 111°31'E], Lanjak-Entimau P.F, c. 2100 ft, Chai S34034
(E, K). Bukit Ubah Ribu, Ulu Sungai Kaup, Lubok Antu [1°03'N 111°50'E], c. 200
ft, Chai S33761 (E).
Cyrtandra penduliflora var. grossipilosa Hilliard & B.L.Burtt var. nov. a planta
Cyrtandra sect. Pleuroschisma 43
typica pilis longis grossis patentibus in caulibus petiolis et subtus in costis (nec pilis
tenuibus valde appressis) filamentorum apicibus et antherarum connectivo dense
glanduloso-puberulo (nec glabro vel fere glabro) differt.
Type: Sarawak, ascent to Gunung Mulu [4°O1'N 114°52'E], 13 vi 1962, Burtt &
Woods B2060 (holo. E).
Unbranched herb, stem up to | m long, 15 mm diam., base often decumbent, rooting,
young parts villous, hairs coarse, spreading. Leaves opposite, strongly anisophyllous,
few at apex of stem, reduced leaves stipule-like, largest developed leaves 190-370 x
85-160 mm, elliptic, apex acute to acuminate, base cuneate, very narrowly decurrent,
margins entire to irregularly toothed, lateral veins 12—18 on each side of midrib, upper
surface glabrous at maturity, coarsely pitted, lower surface with coarse spreading
hairs scattered among short appressed ones on the midrib, similar appressed hairs on
the blade; petiole 100—150 mm long, hairy as midrib. /nflorescence as in the typical
plant, peduncle c. 45—240 mm long. Bracts c. 24-45 x 4-13 mm, finely pubescent.
Calyx lobes c. 6-9 mm long, outside puberulous, inside minutely gland-dotted. Corolla
(only one flower seen) 41 mm long, white to pale yellow with 2 orange-yellow bars in
throat, outside puberulous, mouth glandular-puberulous inside, tube 27mm long,
cylindric in lower part, expanded above, upper lobes c. 8 x 8 mm, lower lip c. 14 x 22
mm, median lobe 9 x 8 mm, all lobes suborbicular. Stamens inserted 22 mm above
base of tube, filaments c. 5 mm long, glandular-puberulous in upper part, strongly
twisted post anthesis, anthers 3 x 2 mm, cohering face to face by a small apiculus,
connective densely glandular-puberulous. Disc 1 x 1.5 mm, cupular. Ovary 6 x 1.2
mm, very minutely pustulate, coma of hairs at apex. Style 13 mm long, glandular-
pubescent. Stigmatic lobes 3 x 2.5 mm, clavate. Fruit c. 25-35 x 5 m, pericarp
smooth. Seeds c. 0.4 x 0.2 mm, testa red-brown.
Notes: This plant differs from typical Cyrtandra penduliflora in the coarse spreading
hairs on the stem and along the midrib on the undersurface of the leaf (versus fine
strongly appressed ones). The tertiary venation also differs subtly : in the typical
plant, the tertiary veins are not clearly visible on the lower surface of the leaf and they
are more or less pinnate in arrangement; in the variety they are clearly visible,
subscalariform, with a fine reticulum of lesser veins between them. The leaves closely
resemble those of C. sarawakensis, distinguished at once by its compact inflorescence.
The label on Sands 5390 (cited below) described the leaves as ‘paler beneath with
reddish-brown veins’; Church 2210 ‘whitish-green beneath with purple veins distinctly
raised’. There is also a difference in the shape of the leaves, those of the typical plant
being broader in relation to length than in the variety, ratio of length to breadth being
1.2—1.6:1 versus 1.7—2.5:1; in absolute terms c.150—200 mm versus 85—160 mm
(largest leaves).
44 Gard. Bull. Singapore 55 (2003)
Anatomically there is some difference in the thickenings of the tracheoids in
the hypodermis: parallel bars in the typical plant, reticulate thickenings in the variety
often mixed with others with parallel bars.
The anther-connective in the typical plant is glabrous or nearly so, that in the
variety densely glandular-puberulous.
From what is known of the distribution of the species, the variety tends to be
more northern and eastern than the typical plant though the few collections from
Kalimantan indicate close proximity.
Other specimens examined: Brunei. Temburong river at Wong Nguan rapids, 4°31'N
L15°15'E, 120m alt., Coode 6649 (E). Temburong distr., Sungai Belalong, 4°30'N
115°10°'E, c. 100 m, Argent et al. 9/1 (E). Slopes above Temburong headwaters NE
of Gunung Retak, 4°22'N 115°17'E, Sands 5309 (E). Tutong, Ladan Hills Forest
Reserve, upstream from Belabau on E bank of Tutong river, 4°26'N 114°46'E, 20 m,
Coode 6409 (E). Sungai Tokat, a branch of Sungai Batu-Apoi, 50-100 m, Hotta
13759 (E). Sarawak. Limbang [4°45'N 115°E], Ulu Medamit [4°27'N 114°55'E],
Sungai Ensungei, Tanjong Long Amok, George et al. $4¢2368 (E). Gunong Api, Ulu
Melinau, Tutoh, 4°07'N 115°15'E, 400 ft, Anderson S31784 (E). Melinau Gorge
pathway, c. 4°05'N 114°50'E, Burtt & Woods B2239 (E). Gunong Mulu National
Park, Gua Rusa, c. 4°02'N 114°50'E, Argent et al. 809 (E). Ulu Belaga, Sungei
Semawat, 3°N 113°54'E, c. 250 m, Hansen 623 (E). Dulit Range [c.3°16'N 114°14'E],
Sungai Sirui, 200 m, Awa & Yii $46629 (E). Valley of Sungai Keyan Linau-Balui
divide, Sungai Nawai, c. 2°26'N 114°10'E, c. 2600 ft, Burtt B1 1456 (E). South of
Punan Lusong, c. 2°28'N 114°12'E, Burtt B] 1294 (E). Kalimantan. Sintang, Bukit
Baka National Park, Sungai Ella and environs, 0°38'S 112°17'E, 320m, Church 280
(A, E, HBG). Serawai, Uut Labang, 750 m, 0°36'6"S 112°38'56'E, Church 2210 (E).
2. Cyrtandra sarawakensis C.B. Clarke, DC., Monog. phan. 5:209 (1883); B.L.
Burtt, Notes Roy. Bot. Gard. Edinb. 30:38 (1970).
Lectotype (Burtt 1970): Sarawak, Kuching, July 1865, Beccari 153 (lecto FI,
isolecto K, P).
Unbranched herb, stem c. 110—230 x 7-20 mm, erect or decumbent and rooting,
young parts densely appressed-pubescent, hairs c. 2 mm long. Leaves opposite, strongly
anisophyllous, c. 3-8 forming a fan at apex of stem, reduced leaves stipule-like, c. 30
x 3 mm, lanceolate, acuminate, closely appressed-pubescent; largest developed leaves
c. 225-300 x 57-125 mm, elliptic, apex very acute, base cuneate, very narrowly
decurrent, margins subentire to serrate, lateral veins 16—20 on each side of midrib,
tertiary veins subscalariform, upper surface glabrous at maturity, coarsely pitted, lower
surface appressed-pubescent on blade, long (to 3mm) delicate appressed hairs on
midrib and lateral veins (tawny or purple in life); petiole c. 80-150 mm, hairy as
Cyrtandra sect. Pleuroschisma 45
midrib. /nflorescence a several-flowered congested cyme, borne mainly in axils of
fallen leaves, peduncie c. 5—30 mm long, stout, puberulous. Bracts persistent,
c. 20-35 x 7-20 mm, lanceolate, acuminate, margins entire or toothed, strongly veined,
both surfaces appressed-pubescent; bracteoles similar but smaller. Pedicels c.10—12 mm,
puberulous. Calyx 5-lobed almost to base, tube c. 0.5 mm, lobes subequal, 4.5—8 x
1—1.5 mm, narrowly deltoid, patent-puberulous outside, inside minutely gland-dotted.
Corolla to c. 44 mm long, palest yellow, 2 orange-yellow bars on palate, outside
puberulous, hairs acute, tube c.27 mm long, narrowly cylindric below, abruptly
expanded above, two upper lobes c. 9 x 9 mm, lower lip 3-lobed, c.14 x 24 mm,
median lobe c. 8 x 8 mm, all lobes suborbicular, glandular-puberulous inside, with
patch of longer hairs adjacent to sinus of upper lobes. Stamens inserted 20-25 mm
above base of tube, c.6mm long, strongly twisted after anthesis, minute glandular
hairs fringing connective; lateral staminodes c. 2 mm long, posticous staminode c. 1 mm.
Disc c.2x 2mm, cupular. Ovary c.7 x 1.4mm, minutely gland-dotted, coma of hairs
at apex. Stigmatic lobes c. 3.5 x 1.2 mm, spathulate, conspicuous, stigmatic papillae
relatively long. Fruit c. 20-25 x 4-5 mm, pericarp smooth. Seeds c. 0.25 x 0.2 mm,
testa bright red-brown, reticulate.
Notes: Cyrtandra sarawakensis was originally collected by Beccari at Kuching.
Beccari had a hut on Mt. Matang, not far from Kuching, and this is possibly the type
locality; Burtt & Woods B1947 (cited below) precisely matches the isolectotype
specimen in Paris. The other collections seen came from the Semengoh Forest Reserve
a few miles south of Kuching. Many species of Cyrtandra seem to have a very limited
distribution and this may be the case here. The distinguishing features of C.
sarawakensis are the many lateral veins, tertiary veins subscalariform and clearly
visible especially on lower surface, hairs on veins (lower surface) strongly appressed,
peduncles c. 5—30 mm long, bracts broad, strongly and conspicuously veined. It is a
plant mainly of the forest floor.
Tracheoids with parallel bars occur in the 1—2-layered hypodermis of the
leaf, and there are no sclereids in the spongy mesophyll. Stomatal turrets are well
developed.
Other specimens examined: Sarawak. Mt. Matang [1°36'N 110°11'E], Burtt & Woods
B1947 (E); Matang, Ulu Sungai Rayu, Lee $54099 (E); Semengoh Forest Reserve,
12 miles S of Kuching [1°28'N 110°22'E], Burtt & Martin B4712 (E); Semengoh
Forest Reserve, Burtt & Woods B2487 (E).
Cyrtandra sarawakensis var. longipilosa Hilliard & B.L.Burtt var. nov. a planta
typica primum pilis in caule et costa grossis et patentibus (nec tenuibus et valde
appressis) et calycis lobis plerumque longioribus 4.5—8 mm (nec 2.54 mm) et apicibus
foliorum longe acuminatis (nec acutis) differt.
46 Gard. Bull. Singapore 55 (2003)
Type: Sarawak [c.2°N 114°10'E, Hose Mts.], Melinau Community Forest, near base
camp at Nanga Tunoh, Burtt & Martin B4772 (holo. E).
Notes: The variety has been collected north and east of Kuching. It resembles
Cyrtandra sarawakensis in nearly all features, including habit, number of lateral veins
in leaves, subscalariform tertiary venation, bracts and other floral characters, but it
differs in its long-acuminate (not merely acute) leaf tips, hairs on stem, petiole and
midrib on under surface of leaves being coarse and spreading (not fine and closely
appressed) and possibly also in length of calyx lobes, 2.54 mm (not 4.5—8 mm).
The coarse spreading hairs are brightly coloured in life, and in the dried state
the colour often persists in the cross walls of the hairs; the hairs on C. sarawakensis
may also be coloured, but, being so much finer, the colour is more difficult to see in
dried specimens. The occurrence of a variety with long coarse spreading hairs as
opposed to appressed ones is paralleled in C. penduliflora, a species with tertiary
venation indistinguishable from that of C. sarawakensis; C. penduliflora differs
markedly in its lax inflorescence, and many specimens have fewer lateral veins in the
leaves.
Other specimens examined: Sarawak. Tatau, path to Bukit Buan, 300 ft. [c.2°56'N
112°55'E], Purseglove 5469 (E). Tatau, Ulu Anap [c. 2°56'N 112°49'E], Mokhtar
S448]5 (E). Bukit Kana [2°42'N 112°54'E], 600-850 m, Hirano & Hotta 1299 (KYO);
Bukit Kana, 50-150 m, Hirano & Hotta 15480 (E). Ulu Anap, Bukit Mersing [2°30'N
113°06'E], c.1000 m, Ashton $17679 (E). Hose Mts., Bukit Temedu [2°24'N 113°41'E],
c. 1100 m, Ashton $19007 (E). Pelagus Rapids on Rajang River [‘Rejang’], also on
Bukit Raya nearby, c. 2°10'N 113°E, Burtt & Woods B2544 (E). Bukit Raya, Smith §
27719 (E). SE end Hose Mts., Bukit Nibong, c. 2°6'W 113°42'E, Burtt & Martin
B4845 (E). Sungai Bena, a tributary of Sungai Sut, c. 1°55'N 113°5'E, Burtt B2609
(E). Lundu distr., Gunung Gading [1°44'N 109°50'E], Burtt & Woods B2685 (E).
Bukit Lumut, Ulu Amau [1°27'N 112°04'E], c. 950 m, Ashton $21264 (E). Bukit
Goram, Ulu Sungai Kapit [1°34'N 112°45'E], Chai S36172 (E).
3. Cyrtandra coacta Hilliard & B.L.Burtt, sp. nov. a C. sarawakense C.B.Clarke
venis lateralibus foliorum 9-15 (nec 15-20), venis tertiariis plus minusve pinnatis
(nec subscalariformibus) pilis in bracteis et in foliorum pagina inferiore papyraceo-
coactis (nec pilis singulatis manifestis) distinguenda.
Type: Sabah, Ranau distr., road from Lohan to Mamut Copper Mine near Tank
47,6°O1'N 116°41'E, 1100 m, 9 vii 1984, Beaman 10656 (holo E).
Cyrtandra sect. Pleuroschisma 47
Unbranched herb to c. 800 mm tall, stem horizonal at base then curving upwards,
clad in a papery indumentum of matted hairs, leafy and floriferous on erect part.
Leaves opposite, strongly anisophyllous, reduced leaf stipule-like, c. 25 x 2 mm,
lanceolate, enveloped in papery indumentum; developed leaf c.180—360 x 60—110(—-165) mm,
elliptic to broadly elliptic, apex acute to almost acuminate, base cuneate, very shortly
decurrent, margins entire to obscurely or distinctly remotely serrate, lateral veins
9-15 on each side of midrib, tertiary veins more or less pinnate, upper surface glabrous,
finely and closely pitted, lower surface papery, any loose hairs very delicate, toc.3mm
long; petiole 75—135 mm, enveloped in papery indumentum. /nflorescence a several-
flowered, very congested, axillary cyme, almost sessile. Bracts persistent, c. 35-40 x
10-16 mm, lanceolate, strongly veined, enveloped in papery indumentum, bracteoles
similar but smaller. Pedicels c.5 mm. Calyx 5-lobed almost to base, lobes subequal,
c. 3.5 x 1 mm, narrowly deltoid, tube c. 0.5 mm, outer surface with papery indumentum,
inside minutely gland-dotted. Corolla c. 40 mm long, white or cream with 2 orange-yellow
bars in throat below the two sinuses of lower lip, outside minutely puberulous, hairs
acute, tube c. 25.5 mm, cylindric below for c.12mm then abruptly expanded, two
upper lobes c. 9 x 6.5 mm, lower lip 3-lobed, c.14.5 x 12 mm, median lobe c. 9 x 7.5
mm, all lobes more or less suborbicular, glandular-puberulous inside and down tube
to insertion of filaments. Stamens inserted c.19mm above base of tube, filaments c. 5
mm, strongly coiled post anthesis, glandular-puberulous at apex, anthers c. 2 x 1.5 mm,
cohering face to face by a prominent ligature, connective densely glandular-puberulous;
lateral staminodes c. 2 mm, posticous staminode c.lmm. Disc 1.5 x 2 mm, cupular.
Ovary c. 8 xX 2 mm, minutely pustulate, coma of hairs at apex. Style c. 15 mm,
glandular-puberulous. Stigmatic lobes c. 3 x 1.5 mm, clavate, conspicuous, stigmatic
papillae relatively long. Fruitc. 20 x 6mm (Mendum & Lamb 234A), pericarp smooth.
Seeds c. 0.5 x 0.25 mm, testa red-brown.
Notes: This species has the aspect of Cyrtandra sarawakensis but is at once
distinguished by the indumentum on the vegetative parts: on stem, leaves (including
petioles), bracts, bracteoles, pedicels and calyx, the hairs are matted together (coactum
meaning felt), giving a papery look to the surface; in C. sarawakensis, hairs on the
corresponding parts are appressed, the individual hairs all separate. The very striking
difference is easily seen along the midrib on the lower surface where the matted hairs
of C. coacta form a papery-looking skin in sharp contrast to the strongly appressed
but distinctly separate hairs in typical C. sarawakensis and the long coarse spreading
hairs in var. Jongipilosa. The indumentum of C. hoseana and C. angustielliptica is
not unlike that of C. coacta but both species differ, inter alia, in their narrow bracts
lacking conspicuous venation.
In C. coacta, there are 9—15 lateral veins on each side of the midrib, in C.
sarawakensis 15—20, while the tertiary veins are more or less pinnate in C. coacta,
48 Gard. Bull. Singapore 55 (2003)
subscalariform in C. sarawakensis. Furthermore, the leaves differ in anatomical detail:
the tracheoids in the hypodermis of C. coacta have reticulate bars and there are no
stomatal turrets; in C. sarawakensis the tracheoids have parallel bars and there are
prominent stomatal turrets.
The stamens may differ too, but too few flowers have been seen to be certain:
in C. coacta, the connective in the anther and the top of the filament are densely
glandular-puberulous; in C. sarawakensis both are glabrous or very minutely glandular.
Cyrtandra coacta is known mainly from Sabah and the northern part of
Sarawak (Yii et al. S5/691 came from the upper reaches of the Baleh River, in east
central Sarawak), while C. sarawakensis is found in the southern part.
Other specimens examined: Sabah. Sinsuron road, 850—900 m, 5°40'N 116°22'E,
Mendum & Lamb 23 (E), Sinsuron road, Mendum & Lamb 23A (E); Lahad Datu
distr., Research Centre, Ulu Sungai Segama, 200m, Argent et al. SAN 108203 (E).
Mt. Kinabalu, eastern shoulder, 6°05'N 116°36-40'E, 3400 ft, Chew et al. 963 (E, K),
Chew et al. 647 (E, K). Sarawak. Seventh division, Ulu Belaga, Sungai Semawat,
3°N 113°54'E, c. 250 m, Hansen 715 (E). NE slope Bukit Kama [2°42'N 112°54'E],
c. 450 m, Hirano & Hotta 1239 (E, KYO). Ulu Sungai Balan, Bukit Batu Tiban, Yii
et al. SS1691 (E).
4. Cyrtandra tunohica Hilliard & B.L.Burtt sp.nov. a C. seganica Hilliard & B.L.Burtt
foliis minoribus (maxime 170—200 mm, nec 300 mm longis) et pro rata angustioribus
(4—-5.7:1, nec 2.5-4.5:1), bracteis plerumque majoribus (20-30 x 4-7.5 mm prope
basin, nec 20 x 4 mm), costa et venis lateralibus praecipue in pagina inferiore visibilibus
(nec costa vix, lateralibus haud visibilibus), calycis lobis 2-4 x 1 mm (nec 4.5—-6 x1.5
mm) distinguenda.
Type: Sarawak, Kapit distr., Sungai Melinau [Hose Mts.], Nanga Tunoh,
c.2°3'40"N 113°38'20"E, c.450ft, Burtt B12664 (holo E).
Unbranched herb, stem c. 5—6 mm in diam. near apex, height unknown, base decumbent
and rooting, erect part toc. 150 mm long, upper part finely pubescent, soon glabrescent.
Leaves opposite, crowded near apex of stem, slightly to strongly anisophyllous even
on one stem, highly reduced leaves, when present, to c. 65 x 2mm, almost filiform,
major developed leaves 170-200 x 35-50 mm, narrowly elliptic, apex acuminate,
base narrowly cuneate, very narrowly and shortly decurrent, margins obscurely serrate
to crenulate, lateral veins 13-15 (as few as 10 in minor leaves), tertiary venation
subscalariform, upper surface glabrous at maturity, pitted, lower surface thinly hairy
on blade, more densely so on veins, all hairs to c. 1 mm long, appressed; petiole 50—
70 mm long, hairy as midrib. /nflorescence: few-flowered congested cymes, axillary,
Cyrtandra sect. Pleuroschisma 49
extending from near apex of stem to well down below the leaves, almost sessile.
Bracts persistent, 20-30 x 4—-7.5 mm, lanceolate, long-acuminate, both surfaces
puberulous, midrib and lateral veins clearly visible especially on lower surface;
bracteoles similar but much smaller. Pedicels c. 1-8 mm, puberulous. Calyx 5-lobed
almost to base, tube c. 0.5 mm, lobes subequal, 2-4 x 1 mm, narrowly deltoid, outside
and margins closely appressed-pubescent, inside minutely gland-dotted. Corolla c.
30 mm long, cream with yellow marks on palate, outside puberulous, hairs acute,
tube c. 20 mm long, lower half narrowly cylindric, upper part expanded, two upper
lobes c. 5 x 3.5 mm, lower lip 3-lobed, median lobe c. 5 x 3.5 mm, all lobes subrotund,
glandular-puberulous inside, hairs extending down into throat. Stamens inserted c.14
mm above base of tube, filaments c. 4 mm long, strongly twisted, anthers 2 x 1.2 mm,
cohering face to face by a small ligature, both anthers and filaments glabrous; lateral
staminodes c. 2 mm, posticous staminode c. 0.5 mm. Disc 1—1.2 x 1.2—1.8 mm,
cupular. Ovary c. 3.5—7 x 0.8—2 mm, very minutely gland-dotted, crowned with
coma of relatively long hairs. Style 5-6 mm long, puberulous. Stigmatic lobes
3—4 x 1-2 mm, clavate. Fruit 10-18 x 4-6 mm, pericarp smooth, deeply grooved
longitudinally. Seeds 0.3-0.5 x 0.2—0.3 mm, testa red-brown.
Notes: The outstanding feature of Cyrtandra tunohica as we know it is variation in
the degree of anisophylly; in the rest of the species in this section of Cyrtandra, the
minor leaf is always reduced to a stipule-like outgrowth (with the caveat that several
species are ill-known). In C. tunohica, the size of the leaves in each pair ranges from
subequal (when the stem is much more leafy than is usual in this section as a whole)
through degrees of reduction culminating in an extraordinarily long, almost filiform,
stipule-like leaf; this occurs on a single stem. As the species is known from only one
locality, the taxonomic value of this character remains to be assessed. The plants
(very few) were found on the steep shady earth bank of a stream gulley, and were seen
only once, despite further fieldwork in the area on this and other occasions.
In its relatively narrow leaves, C. tunohica resembles both C. seganica and
C. angustielliptica; it is easily distinguished from both by its bracts, 4-7.5 mm broad
near the base (not 1.8—4 mm) and with both midrib and lateral veins raised on the
lower surface and thus clearly visible (in contrast to nearly or quite invisible). It is
further distinguished from C. seganica by its leaves, the largest 170-200 mm long
(not c. 300 mm) and relatively narrower (ratio of length to breadth 4—5.7:1 versus
2.5—4.5:1), short (to c.1mm long) appressed hairs along the midrib on the
lowersurface (not long coarse spreading hairs), and c.13—15 (not 18) lateral veins on
each side of the midrib.
Not only do the bracts differ from those of C. angustielliptica, but so do the
leaves in several details: lateral veins 13-15 in major leaves (not 7-10), hairs on
undersurface of leaves up to c.1mm long, sparse on the blade (in contrast to silky-villous,
hairs c. 2 mm long, the blade eventually glabrous, hairs persistent on midrib), tertiary
veins Clearly visible (not invisible or very nearly so).
50 Gard. Bull. Singapore 55 (2003)
The leaf of C. tunohica has tracheoids with reticulate thickening in the
hypodermis and lacks sclereids in the mesophyll; C. seganica and C. angustielliptica
have similar tracheoids but differ in having polymorphic sclereids in the mesophyll.
Other specimen examined: Sarawak. Kapit distr., Sungai Melinau [Hose Mts.], Nanga
Tunoh, c. 2°3'N 113°38'E, Burtt B12665 (E).
5. Cyrtandra sp. nov.
Unbranched herb, stem very stout, c. 12 mm diam., c. 300-600 mm tall, young parts
villous, lower parts glabrous. Leaves opposite, few crowded near apex of stem, strongly
anisophyllous, reduced leaves stipule-like, c. 40 x 2 mm, narrowly linear-lanceolate,
villous; largest developed leaves c. 195-260 x 80-90 mm, elliptic or oblanceolate,
apex abruptly acuminate, base narrowly cuneate, very narrowly decurrent, margins
entire, lateral veins 16—18, tertiary veins finely reticulate, upper surface glabrous,
coarsely pitted (at maturity; densely pubescent when juvenile), lower surface densely
pubescent, veins pilose; petiole c. 30-70 mm long, villous. /nflorescence a many-flowered,
highly congested dichasial cyme, axillary, forming bracteate clusters borne on the
leafless part of the stem; peduncle very stout, to 2 mm long. Bracts persistent, primary
pair c.20x7mm, lanceolate, acute, midrib and lateral veins prominent, outside densely
pubescent, inside hairs mainly near apex and base; bracteoles similar but narrower, a
pair subtending each pedicel. Pedicels c.5 mm, puberulous. Calyx 5-lobed almost to
base, tube c. 0.5 mm, lobes c. 4 x 1 mm, outside puberulous, hairs patent, inside
minutely gland-dotted. Corolla c. 30 mm long, white, 2 orange-yellow bars on palate,
outside puberulous, hairs acute, tube c. 20 mm long, lower part narrowly cylindric,
upper part expanded, two upper lobes c. 5 x 3.5 mm, lower lip 3-lobed, median lobe
c.5 x 4mm, all lobes subrotund, glandular-puberulous on lower lip, hairs extending
down floor of tube, a median tuft of eglandular hairs at base of each upper lobe, a few
glandular hairs towards outer margins. Stamens inserted c.17 mm above base of tube,
filaments c. 4-5 mm long, twisted once near base, anthers 2.2 x 1 mm, cohering face
to face by a conspicuous ligature, connective minutely glandular, glands extending
briefly down back of filaments; lateral staminodes c. 2 mm, apex strongly hooked, tip
globular, pale yellow (as anthers), posticous staminode c.1 mm. Disc 1.8 x 2 mm,
almost cupular (deeply notched on one side). Ovary 7 x 2 mm, glabrous except for
conspicuous coma. Style 10 mm (at anthesis), pubescent, hairs acute. Stigmatic
lobes c. 2 x 1.5 mm (post anthesis), spathulate, stigmatic papillae conspicuous. Fruit
23 x 6 mm (one seen), pericarp smooth, split longitudinally, only apex and base
cohering. Seeds c. 0.25 x 0.2 mm, testa red-brown.
Notes: This species is allied to Cyrtandra tunohica and C. sarawakensis var.
Cyrtandra sect. Pleuroschisma 34
longipilosa. It is distinguished from C. tunohica by the relatively long patent
indumentum on stems, petioles and backs of leaves (versus hairs short, appressed),
leaves c. 80-90 mm broad (not 35-50 mm) and many- (not few-) flowered
inflorescence. It differs from C. sarawakensis var. longipilosa by its shorter petioles
(30-70 mm, not 80—150 mm) and almost sessile inflorescence. Also, the pubescence
on the undersurface of the leaf blade is dense and patent (not appressed). This is
obvious in Haron $29978 (K), but not so well marked on the duplicate (E); it is also
obvious in Jugah Kudi S23712 (K) where the finely reticulate tertiary venation shows
up well.
The two collections seen are not good: Haron S29978 was obviously pressed
when badly withered; the sheet at Kew is particularly bad, but this is the one with a
nearly complete corolla and several buds, from which the floral details were extracted.
Only an old fruit survives on the sheet in Edinburgh, and the base of a fruit on Jugah
Kudi S23712 appears to have eight loculi! The need for good collections before the
species can be named is obvious.
The leaves have a 1—2-layered hypodermis, and tracheoids with parallel bars
in the hypodermis.
Specimens examined: Sarawak, Anap, Ulu Kakus [c.2°30'N 113°E], Othman Haron
$29978 (E, K); Balingian, Ulu Sungai Arip, Bukit [ju [roughly 2°40'N 112°40'E],
Jugah Kudi §23712 (K, L).
6. Cyrtandra tesselata Hilliard & B.L.Burtt sp. nov. a C. tunohica (etiam folia anguste
lanceolata gerente) caulibus et pagina foliorum inferiore villosis (nec breviter
pubescentibus), petiolo 20—25 mm longo (nec 50—70 mm) villoso (nec pubescente),
bracteis c.15 x 1.8—2 mm venatione invisibili (nec 20-30 x 4-7.5 mm venatione elevata
conspicua), calycis lobis c.6—8 mm longis (nec 2.4 mm) differt.
Type: Sarawak, Bintulu distr., en route (survey route) from Sungai Mah to Sungai
Shinonok, Ulu Sungai Minah, alt. 40-150 m, Hotta 1/4117 (holo KYO).
Herb, probably unbranched, height unknown, stem collected 50 x 10 mm, villous,
hairs 3-4 mm long, leafy throughout, internodes c. 10 mm long. Leaves opposite,
strongly anisophyllous, reduced leaves stipule-like, c. 30 x 3 mm, narrowly
lanceolate,long-acuminate, major mature leaves 150-230 x 26-34 mm, narrowly
elliptic, apex gradually very acute, base narrowly cuneate, margins very obscurely
serrulate, lateral veins 12—14 on each side of midrib, upper surface glabrous at maturity,
pitted, lower surface villous, longest hairs (2-3 mm) on midrib, shorter on lesser
veins, tertiary venation finely reticulate, all veins raised, hairs brownish (dried state)
making venation very conspicuous; petiole 20—25 mm long, villous. Inflorescence a
few-flowered axillary cyme, peduncle c. 4mm, very stout. Bracts c. 15x 1.8—2 mm,
narrowly lanceolate, densely pubescent, venation obscure; bracteoles similar but
52 Gard. Bull. Singapore 55 (2003)
Pedicels c.10mm (in fruit), pubescent. Calyx 5-lobed almost to base, tube c. 0.5 mm,
lobes 6-8 x 1 mm in fruit, narrowly deltoid, outside and margins puberulous, hairs
patent, inside minutely gland-dotted. Corolla not seen, only very young buds present,
puberulous outside, hairs acute, lobes minutely glandular inside (tube scarcely
developed). Stamens: filaments scarcely developed, anthers 2 x 0.8 mm, minutely
glandular on margins of connective. Disc cupular. Ovary and style very young, style
pubescent. Fruit c. 35 x 5 mm, pericarp smooth, traces of coma still visible at apex.
Seeds c. 0.25 x 0.2 mm, testa bright red-brown.
Notes: This species is so distinctive that lack of flowers should not preclude its formal
recognition. The leaves are closely set down the 150 mm length of the piece of stem
on the herbarium sheet, in contrast to most of its allies where the leaves appear to be
quickly deciduous leaving a few, forming a fan, at the apex of the stem. Unfortunately,
the collector gave no notes either on habit or habitat of his plant. The major leaves are
short-petioled, narrowly elliptic, and at maturity glabrous above but villous below,
with midrib, lateral and tertiary veins raised (the latter forming a fine reticulum) and
conspicuous, partly owing to the dark hairs clothing them. Similar tertiary venation
has not been seen in any other species in this section.
Cyrtandra tesselata is diagnosed above against C. tunohica, which has
similarly narrowly lanceolate leaves, but the indumentum on stem, petioles and leaf
undersurface in C. tesse/ata is villous (not merely shortly pubescent), the petioles are
remarkably short (20-25 mm versus 50-70 mm), the bracts are c. 15 x 1-2 mm,
venation invisible (not 20-30 x 4—7.5 mm, venation raised and conspicuous), calyx
lobes c. 6-8 mm long (not 24 mm). In sharp contrast to that of C. tesselata, the
tertiary venation in the leaves of C. tunohica is rather indistinct, and tends to be
subscalariform.
The leaf anatomy of C. tesselata is unlike that of C. tunohica: vertically long
tracheoids with parallel bars in the hypodermis, polymorphic sclereids in the
mesophyll; the tracheoids in C. tunohica have reticulate bars and there are no sclereids
in the mesophyll.
7. Cyrtandra seganica Hillard & B.L.Burtt, sp. nov. a C. sarawakense C.B.Clarke
bracteis tantum c. 4 mm latis (nec 7-20 mm) sine venatione conspicua (nec costa et
venis lateraliibus prominentibus) et folia angustiora prorata longitudinis (5.5:1 nec
2.54.5:1) facile distinguenda.
Type: Sarawak, Bintulu distr., Ulu Segan [c. 2°30'N 113°E], rocky stream bank, 750
ft., Jlias Paie §27215 (holo E).
Cyrtandra sect. Pleuroschisma 53
Unbranched herb, height unknown, stem c. 8 mm diam. near apex, apex densely
pubescent, rest of stem quickly glabrous. Leaves opposite, strongly anisophyllous,
few at apex of stem, reduced leaves stipule-like, c. 25 x 3 mm, lanceolate, acuminate,
densely pubescent, hairs spreading; developed leaves c. 300 x 55—60 mm, elliptic,
acute, base narrowly cuneate, very narrowly decurrent for c. 35 mm, margins entire,
lateral veins c. 18 on each side of midrib, tertiary venation subscalariform, upper
surface glabrous, pitted, lower surface with very small fine strongly appressed hairs
on blade, coarse much longer (c. 1 mm) hairs on midrib and lateral veins, probably
appressed and probably purplish in life (specimens well glued); petiole c. 55-70 mm
long, hairy as midrib. /nflorescence: few-flowered congested cymes borne mainly in
axils of fallen leaves, peduncle c. 4-8 mm long, very stout, puberulous. Bracts
persistent, c. 20 x 4mm, narrowly lanceolate, acuminate, margins entire, both surfaces
puberulous, hairs spreading, midrib scarcely visible; bracteoles similar but smaller.
Pedicels c. 8-10 mm. Calyx lobed nearly to base, lobes subequal, 4.5—6 x 1.5 mm,
narrowly deltoid, tube c. 0.5 mm, outside and margins puberulous, inside minutely
gland-dotted. Corolla not seen. Fruit c. 18—36 x 4—5.5 mm, pericarp smooth. Seeds
immature.
Notes: Cyrtandra seganica is currently known with certainty only from the type
collection. In the venation of the leaves, including the subscalariform arrangement of
the tertiary veins, it resembles C. sarawakensis, but the leaves are narrower in relation
to their length (ratio of length to breadth 5.5:1 versus 2.5-4.5:1). There is a striking
difference in the bracts, only c. 4 mm broad near the base and with the midrib but
faintly visible in C. seganica, in contrast to bracts c. 7-20 mm broad with prominent
venation in C. sarawakensis. The leaves are similar anatomically in having tracheoids
with parallel bars in the hypodermis, but differ in the presence of many polymorphic
sclereids in the mesophyll of C. seganica whereas the leaves of C. sarawakensis lack
sclereids.
Two collections from further south and west resemble C. seganica in
leaf-venation and in narrow bracts. They appear to differ in the shape of the leaves
(oblanceolate) and also in the ratio of length to breadth (3.25—-4.6:1), this being within
the range of C. sarawakensis. The leaves also appear to differ anatomically, in that
the tracheoids have reticulate bars, not parallel bars; polymorphic sclereids occur in
the mesophyll. These two collections come from Sarawak from Datu Permanent Forest
[Bukit Datu? at 1°29'N 110°50'E], 420 m alt., on the ridge between two streams,
Lee S§41904 (E) and 17" mile Bau/Lundu road, Gunong Undan (1° 27'N 109° 59'B),
stream bank, 50m, Yii S45942 (E).
Much more needs to be known about all three plants described here.
54 Gard. Bull. Singapore 55 (2003)
8. Cyrtandra linauana B.L.Burtt, species nova ex affinitate C. sarawakensis
C.B.Clarke et C. hoseanae B.L.Burtt, ab ambabus nervis lateralibus foliorum 7-8
(nec 12—20) et calycis lobis 10-15 mm longis (nec usque ad 8 mm) differt.
Type: Sarawak, Belaga distr., hill just N of Long Linau [c.2°40'N 114°E], Burtt 11478
(holo E; iso BO, L, SAR).
Herb, stem simple, c.100 mm tall, 8 mm diam., puberulous, hairs very closely
appressed. Leaves opposite, strongly anisophyllous, 5 pairs crowded near apex of
stem, lower ones fallen, reduced leaf stipule-like, c. 35 x 5 mm, lanceolate, long-acuminate;
developed leaf c. 210-300 x 82-83 mm, elliptic, somewhat falcate and eccentric,
apex rather abruptly acuminate, base cuneate, very shortly and narrowly decurrent,
margins entire, lateral veins 7-8 on each side of midrib, tertiary venation coarsely
reticulate, upper surface mottled silver (living material), glabrous, pitted, lower surface
with blade glabrous, midrib and lateral veins strongly appressed-pubescent, hairs very
inconspicuous; petiole 70-80 mm, thickly appressed-pubescent. J/nflorescence an
axillary dichasial cyme, flowers few, branches of cyme very short (3-4 mm), bearing
only 4 flowers (apical bud on each branch suppressed), peduncle c. 5-8 mm, very
stout. Bracts persistent, c. 20 x 2 mm, narrowly lanceolate, acuminate, both surfaces
strongly appressed-pubescent, bracteoles similar but smaller, a pair subtending each
pedicel. Pedicels c.20 mm, appressed-pubescent. Calyx 5-lobed almost to base, tube
c.0.5 mm, lobes 10—15 mm, narrowly linear-lanceolate, outside appressed-pubescent,
inside minutely gland-dotted. Corolla 26-30 mm long, white, throat yellow shading
to orange, outside minutely puberulous, hairs acute, red, tube 17—18 mm long, cylindric
in lower third, abruptly expanded above, two upper lobes 7—8 x 5 mm, lower lip
3-lobed, c. 9-12 mm long, median lobe 7—8 x 5 mm, all lobes oblong-elliptic, glandular-
puberulous inside. Stamens inserted 8.5 mm above base of tube, filaments 2.5 mm,
glabrous, anthers 2.2x 1.2mm, cohering face to face by a prominent ligature, connective
fringed with stout red hairs; lateral staminodes 3 mm, posticous staminode 1.2 mm.
Disc 1.5 x 2 mm, cupular but deeply excavated on one side. Ovary 5 x | mm, minutely
pustulate, coma of hairs at apex. Style 7mm, glandular-puberulous. Stigmatic lobes
4x 1.8 mm, clavate, conspicuous. Fruit 24 x 3.8 mm, pericarp smooth.
Notes: Cyrtandra linauana resembles both C. sarawakensis and C. hoseana in the
size, shape and stance of its leaves but differs from both in having only 7-8 lateral
veins on each side of the midrib (not 12—20). The tertiary venation also differs: in C.
linauana it is rather indistinct and very coarsely reticulate though the veinlets may be
more subscalariform towards the margins; in C. sarawakensis the veins are prominent
and closely subscalariform; in C. hoseana they are very nearly invisible.
Cyrtandra sect. Pleuroschisma 55
The tracheoids in the hypodermis have reticulate bars as in C. hoseana but there are
no sclereids in the spongy mesophyll. In C. sarawakensis the tracheoidal bars are
parallel and, as in C. linauana, there are no sclereids in the spongy mesophyll; C.
hoseana has polymorphic sclereids.
The inflorescences of all three species differ: in C. /inauana, the bracts are
only c. 2 mm broad and the venation is not or scarcely visible; pedicels c. 20 mm
long, making the inflorescence rather lax. In C. sarawakensis, the bracts are very
conspicuous, being broad, venation prominent, pedicels c. 9-13 mm long, inflorescence
congested. In C. hoseana, the bracts are c. 34 mm broad, venation inconspicuous,
pedicels c. 5-10 mm long, inflorescence congested. Stout red hairs fringe the
connective in C. linauana; in C. sarawakensis and C. hoseana, the connective is
either glabrous or glandular.
Cyrtandra linauana is known with certainty only from the type collection
made at c. 2°40'N 114°E. There is another collection, made at c. 2°48'N 112°59'E,
that is clearly very close to C. linauana, namely Hirano & Hotta 88] (KYO, E) from
Bukit Keyan, Ulu Sungai Kakus, alt. 100-300 m. It differs in several details: leaves
not falcate and with 10 lateral veins, otherwise identical with those of C. linauana,
flowers fascicled in the leaf axils, pedicels c. 15 mm long, calyx lobes 6.5—9 mm.
The most significant difference is the fascicled flowers in contrast to the shortly
pedunculate dichasial cyme of C. linauana, where the long pedicels produce a very
lax inflorescence.
The leaves of C. angustielliptica have a deceptive similarity to those of C.
linauana. That species is, however, distinguished by the short (to c. 1 mm) appressed
and inconspicuous hairs on stem, petiole and midrib of leaves on lower surface (not
2—3 mm long, silky, appressed hairs matted together and easily seen), bracts c. 20 mm
long (not 5—8 mm), connective of anthers fringed with stout red hairs (versus glandular
puberulous ones all over). The inflorescence of C. angustielliptica is a nearly sessile,
very congested, cyme.
9. Cyrtandra hoseana B.L.Burtt, Notes Roy. Bot. Gard. Edinb. 30:33 (1970).
Type: Sarawak, Lambir Hills, c. 4°7'N 113°55'E, sandstone banks, Burtt & Woods
B2365 (holo E, iso SAR).
Unbranched herb, stem 20-150 cm long, c. 8 mm diam. near apex, hanging from
sandstone cliffs and banks, sometimes supported by prop roots, young parts silky
villous, hairs strongly appressed. Leaves opposite, strongly anisophyllous, few
arranged in a fan at stem apex, reduced leaves stipule-like, c. 20 x 10 mm, broadly
lanceolate, appressed silky-villous; largest developed leaves c. 255-320 x 85-95 mm,
elliptic, apex abruptly acute, base cuneate, shortly and very narrowly decurrent, margins
entire to obscurely serrulate, lateral veins 12-15 on each side of midrib, tertiary
venation coarsely reticulate, upper surface glabrous at maturity, coarsely pitted, lower
56 Gard. Bull. Singapore 55 (2003)
surface with long, strongly appressed, silky hairs on midrib and lateral veins, blade
glabrous or almost so at maturity; petiole c. 45-60 mm, hairy as midrib. /nflorescence
an axillary, very congested, cyme, mainly in axils of fallen leaves, few-flowered,
peduncle 2-6 mm long. Bracts c. 15-18 x 34 mm, persistent but inconspicuous,
lanceolate, acuminate, closely appressed-puberulous, only midrib visible on lower
surface. Pedicels c. 5—10 mm long, puberulous. Calyx 5-lobed almost to base, lobes
subequal c. 2.2-3.5 x 1.25 mm, deltoid, outside appressed-pubescent, inside minutely
gland-dotted. Corolla c. 32 mm long, white or creamy white outside, 2 orange-yellow
bars in throat, outside puberulous, hairs acute, red, tube c. 20 mm long, lower half
narrowly cylindric, abruptly expanded above, two upper lobes c. 5 x 7 mm, lower lip
3-lobed, c.12.5 x 20 mm, median lobe c. 6 x 7 mm, all lobes subrotund, glandular-
puberulous inside and for a short way down tube. Stamens inserted c. 16 mm above
base of tube, filaments c. 5 mm, curved and twisted once, minute globular glands at
apex, anthers c. 3 x 1.6 mm, cohering face to face by a small ligature, connective with
a few glandular hairs and globular glands; lateral staminodes c. 1.2 mm, posticous
staminode c.lmm. Disc 1 x 1.8 mm, cupular. Ovary 8 x 1.6 mm, very minutely
gland-dotted, coma of hairs at apex. Style 10mm, glandular-puberulous. Stigmatic
lobes not fully developed, c. 2 mm long, clavate. Fruit c. 27-45 x 5 mm, pericarp
smooth. Seeds c. 0.4 x 0.25 mm, testa red-brown.
Notes: So far, C. hoseana is known with certainty only from the Lambir Hills. Now
that more material is available, it is clear that Burtt & Woods B2212 (E, SAR) from
the Melinau Gorge, which was cited in the original description, is a distinct species,
described here as C. angustielliptica.
It is worth noting again that ‘the youngest leaf is very densely covered with
shining silky hairs and it hangs downwards at the top of the shoot, like a tiny silvery
flag’ (Burtt, 1970). In this species, the tertiary veins are scarcely discernible on the
lower surface of the leaf, but form a coarse reticulum that may just be visible on the
upper surface (compare the tertiary veins of C. sarawakensis).
Tracheoids with reticulate bars occur in the 1—2-layered hypodermis, and a
few polymorphic sclereids in the mesophyll.
Other specimen examined: Sarawak. Lambir National Park, Sungai Lapoh and nearby,
Burtt 11546 (E).
10. Cyrtandra angustielliptica Hilliard & B.L.Burtt sp. nov. a C. hoseana B.L.Burtt
foliis 36-60 mm latis (nec 85-90 mm) apice acuminato (nec abrupte acuto), venis
lateralibus utrinque costae 7-10 (nec 12-15), calycis lobis 5-8 mm longis
(nec 2.2—3.5 mm) distinguenda.
Cyrtandra sect. Pleuroschisma S¥
Type: Sarawak, Sungei Melinau Gorge, c. 4°5'N 114°50'E, sandstone hillock, Burtt
& Woods B2212 (holo E).
Unbranched herb, stem of unknown height, c. 6-7 mm in diam. near apex, young
parts silky-villous, hairs strongly appressed, matted together. Leaves opposite, strongly
anisophyllous, reduced leaves stipule-like, c. 18 x 6 mm, lanceolate, silky-villous as
stem, developed leaves up to c.10, largest 170-275 x 36-60 mm, elliptic, apex
acuminate, base cuneate, margins entire to obscurely serrulate but teeth made
conspicuous by apical tuft of hairs, lateral veins 7—10 on each side of midrib, tertiary
venation obscure, upper surface glabrous at maturity, coarsely pitted, lower surface
with long, strongly appressed, silky hairs on midrib and lateral veins, these also on
blade but this eventually glabrous; petiole c. 40-45 mm, hairy as midrib. /nflorescence
an axillary very congested cyme, flowers few, peduncle c. 2 mm. Bracts c. 5.5—7 x
1.8—2 mm, persistent but inconspicuous, lanceolate, acuminate, both surfaces densely
appressed-pubescent. Pedicels c. 8 mm, puberulous. Calyx 5-lobed almost to base,
tube c. 0.5 mm long, lobes subequal, 5—8 x 1.2 mm (in flower and fruit), deltoid,
outside densely appressed-pubescent, inside minutely gland-dotted. Corolla c. 30-35 mm
long, white or creamy white outside, 2 orange-yellow bars in throat, outside puberulous,
hairs acute, tube c. 22—28 mm, lower half cylindric, upper part expanded, two upper
lobes c. 5 x 5 mm, lower lip 3-lobed, median lobe c. 6 x 7 mm, all lobes subrotund,
glandular-puberulous inside and short way down tube. Stamens inserted c. 16 mm
above base of tube, filaments c. 5 mm, curved and twisted once, glandular-puberulous
at apex, anthers c. 2 x 1.5 mm, cohering face to face by a small ligature, connective
glandular-puberulous; lateral staminodes c. 2 mm, posticous staminode not seen. Disc
c.1.5x 1.8mm, cupular. Ovary c.6x 1.2 mm, very minutely papillose, coma of hairs
at apex. Style c.6 mm elongating to c.10 mm, glandular-puberulous. Stigmatic lobes
c.3x 2mm, clavate. Fruit 25-48 x 5 mm, pericarp smooth. Seeds c. 0.3 x 0.2 mm,
testa red-brown.
Notes: In his original description of Cyrtandra hoseana, Burtt cited Burtt B2212
(above) as that species, but in the herbarium he later segregated it and further collections
as differing in their narrower leaves. Not only are the leaves narrower, they are also
shorter (170—275 x 36-60 mm versus 255-320 x 85-95 mm) and there are significantly
fewer lateral veins than in C. hoseana (7-10 versus 12-15); also, the leaf tips are
acuminate (not abruptly acute). They may differ further in the length of the calyx
lobes (5—8 mm versus 2.2—3.5 mm, but see comments below). As in C. hoseana the
tertiary veins are scarcely visible on the lower surface of the leaf; anatomically the
leaves are similar, having tracheoids with reticulate bars in the hypodermis and
polymorphic sclereids in the mesophyll.
Cyrtandra angustielliptica is known only from the Melinau Gorge (in Gunung
Mulu National Park) at c.100 m above sea level. A plant with leaves exactly like
those of C. angustielliptica in all morphological details has been collected several
58 Gard. Bull. Singapore 55 (2003)
times on the extreme headwaters of the Balleh, at c.1000—1700 ft altitude on a
ridge between Sungai Balang and Sungai Balleh [1°35'N 114°30'E at 1200 ft by
Anderson & Paie S28329 (E); at c.1000 ft by Anderson & Paie 28739 (E); at c.1000 ft
by Anderson $2874] (E) and at 1700 ft by Paie S28409 (E, L)]. However, these
specimens differ anatomically in having no sclereids in the spongy mesophyll (a
character possibly of little taxonomic significance), and the calyx lobes are 2.5—5 mm
long (not 5—8 mm), corolla 20—25 mm (not 30—35 mm), filaments and anthers glabrous
(not glandular), and the disc unilateral (not cupular).
Other specimens examined: Sarawak. Gunung Mulu National Park, Melinau Gorge,
Burtt B8274 (E); Melinau Gorge, c. 100 m, Nielsen 537 (E).
11. Cyrtandra insolita Hilliard & B.L.Burtt sp. nov. nullae arcte affinis; pedunculo
longo axillari bracteis duabus flores breviter pedicellatos 1—3 involucrantibus facile
recognoscenda.
Type: Sarawak, Kapit, Upper Rajang [“Rejang’] River, 1929, Clemens 21236 (holo K).
Shrub (fide collector), 3 stems seen, 170-250 mm long, woody, leafy only at apex,
young parts densely pubescent. Leaves opposite, strongly anisophyllous, reduced
leaves stipule-like, c. 20 x 3 mm at base, linear-lanceolate, villous; developed leaves
few, c.160—190 x 55 mm, elliptic, apex long-acuminate, base cuneate, decurrent down
petiole, margins obscurely serrulate, lateral veins c. 20 on each side of midrib, tertiary
veins invisible, upper surface silky villous at first, soon glabrescent, coarsely pitted,
lower surface densely pubescent, hairs very delicate, c.4mm long on midrib, shorter
on blade; petiole c.10mm long, villous. /nflorescence mainly in axils of fallen leaves,
1—3-flowered, flowers springing from apex of peduncle and enfolded by bracts,
peduncle 15-45 mm long, villous. Bracts 2, persistent, foliaceous, c. 35-45 x 10—22
mm, lanceolate, long-acuminate, 5-nerved, both surfaces villous; bracteoles wanting.
Pedicels 4-5 mm, pubescent. Calyx 5-lobed almost to base, lobes subequal, 2 x 1.5 mm,
deltoid, outside pubescent, inside gland-dotted. Corolla not seen. Anthers (in very
young bud) 1.5 x 1.1 mm, glabrous. Gynoecium very young, but stigma clearly bilobed,
disc cupular. Fruit c. 30 x 5 mm, pericarp smooth (young fruits retain an apical coma).
Seeds c. 0.3 x 0.25 mm.
Notes: Cyrtandra insolita (insolitus = extraordinary) is highly distinctive among other
species in this Section, from which the pair of foliaceous bracts enfolding one to
three flowers at the apex of the long peduncle immediately mark it out. The other
species all have few- to many-flowered cymes. Also, these species are unbranched
herbs of the forest floor; Mrs Clemens described her plant as ‘shrub in thickets’.
Cyrtandra sect. Pleuroschisma aby
12. Cyrtandra sp. nov.
Unbranched herb, stem woody, height unknown, 10 mm diam., pilose, hairs coarse,
spreading, red-purple, as on other vegetative parts. Leaves opposite, strongly
anisophyllous, few at apex of stem, reduced leaves stipule-like, c. 75 x 6 mm at base,
linear-lanceolate, long-acuminate, pilose; largest developed leaves 300-350 x 85-110 mm,
elliptic, apex acuminate, base narrowly cuneate, decurrent, margins finely and remotely
toothed (teeth being extensions of the veins beyond the margins), lateral veins 15—16
on each side of midrib, tertiary veins coarsely reticulate, upper surface glabrous, pitted,
lower surface with short, fine appressed hairs on blade, long coarse spreading hairs
on midrib; petiole c. 45—50 mm long, hairy as midrib. /nflorescence a congested
several-flowered cyme, solitary in axils of fallen leaves, peduncle c. 6-12 mm long,
pubescent. Bracts persistent, c. 18 x 3 mm, lanceolate, acuminate, midrib visible on
lower surface, both surfaces densely pubescent; bracteoles similar but smaller, a pair
subtending each pedicel. Pedice/l c. 20 mm long (in fruit), densely puberulous. Calyx
5-lobed almost to base, tube c. | mm long, lobes subequal, c. 10 x 1.5 mm (in fruit),
narrowly deltoid, outside densely puberulous, inside minutely gland-dotted. Corolla
not seen. Fruit c. 30 x 5.5 mm, pericarp smooth, remains of coma visible at apex.
Seeds not fully ripe.
Notes: This plant needs to be re-collected. Gunung Gaharu lies c. 96 km SE of Kuching
on the road to Serian. The specimen seen is a unicate, in fruit but without flowers. It
much resembles Cyrtandra sarawakensis, particularly var. longipilosa, but is at once
distinguished by the reticulate, not subscalariform, tertiary venation of the leaves,
and by its much narrower bracts.
Specimen examined: Sarawak, Gunung Gaharu, c. 2000 ft., Burtt 2658 (E).
Acknowledgements
We thank the curators of the herbarium at Royal Botanic Gardens, Kew (K), Botany
Dept., Kyoto University (KYO), the Nationaal Herbarium Nederland, Universiteit
Leiden Branch (L) for the invaluable loan of specimens. We are also grateful to
several of our colleagues: Frieda Christie, electron microscope unit; Mary Mendum
(Bates) for art work; Adele Smith for stepping into the breach in a typing crisis; and
Debbie White, photographic unit. We also thank the Regius Keeper, Steve Blackmore,
working facilities at the Garden.
60 Gard. Bull. Singapore 55 (2003)
References
Bokhari, M.H. and B.L. Burtt. 1970. Studies in the Gesneriaceae of the Old World
XXXII: Foliar sclereids in Cyrtandra. Notes Royal Botanic Garden Edinburgh.
30: 11-21.
Burtt, B.L. 1970. Studies in the Gesneriaceae of the Old World XX XIII: Some species
of Cyrtandra chiefly from Borneo. Notes Royal Botanic Garden Edinburgh. 30:
23-42.
Burtt, B.L. 2001. A survey of the genus Cyrtandra (Gesneriaceae). Phytomorphology
Golden Jubilee Issue 2001: Trends in Plant Sciences. pp. 393-404.
Garden’s Bulletin Singapore 55 (2003) 61-63
Begonia peninsulae (Begoniaceae), a Confused Species from
Malesia
RUTH KIEW
Singapore Botanic Gardens, Singapore 259569
Abstract
Begonia peninsulae Irmsch. subsp. peninsulae from Trengganu, Peninsular Malaysia, is
synonymous with B. rajah Ridl. and B. peninsulae subsp. tambelanense Irmsch. is raised to
specific rank as B. tambelanense (Irmsch.) Kiew.
Introduction
Irmscher (1929) recognised two subspecies within his Begonia peninsulae: ssp.
peninsulae from Trengganu and ssp. tambelanense from the Tambelan Islands. These
were both based on a mixture of plants cultivated in the Singapore Botanic Gardens
and the Penang Botanic Garden. The ones collected from the Singapore Botanic
Gardens in Ridley’s time (one from Trengganu; the other from the Tambelan Islands)
have a clear provenance. However, later collections are without provenance and, in
the case of those from Penang, are from batches that include many exotic ornamental
begonias. Indeed, there is no evidence to suggest that they are plants of wild-collected
origin. In addition, these two subspecies are clearly not conspecific as one has a cordate
leaf and the other a peltate one.
Subspecies peninsulae
Irmscher cited one specimen from Trengganu (which was spelt ‘Tringganu’ on the
label), which is without a number or information on the date and collector. The botanical
name is written as ‘Begonia raja’. ‘Ridley’ has been added as the collector in a different
hand after Irmscher’s publication.
This specimen matches those of Begonia rajah Ridl. in characters of the leaf,
inflorescence and fruit. Despite the leaf being exceptionally large (11.5 x 16 cm as
opposed to 5.5—7.5 x 5.5—7.5 cm in wild plants of B. rajah), the leaf base, like B.
rajah, is deeply cordate and, even in the dried state, the variegation is conspicuous
with the veins lighter than the rest of the lamina. This specimen might be one of the
62 Gard. Bull. Singapore 55 (2003)
plants originally collected in Trengganu (which Ridley spelt as Tringganu) or its
progeny that was cultivated in the Botanic Gardens Singapore from where it was
introduced into the ornamental trade in England. B. peninsulae ssp. peninsulae is
therefore reduced to synonomy with B. rajah.
The other specimens of ssp. peninsulae cited by Irmscher are very different
in that they are all peltate (B. rajah never has peltate leaves). These specimens, Burkill
Begonia No. 16 July 1917 (SING), Burkill 6114 31 Aug 1920 (SING), were cultivated
in the Penang Botanic Garden and have no provenance. In leaf size, shape, colour,
surface (‘coppery and bullate’) and margin they are identical to Begonia goegoensis
N.E. Brown from Sumatra, which is one of the few Asian begonias that is easy to
grow under local conditions.
Begonia rajah Rid.
Gard. Chron. II, 16 (1894) 213 & Fig. 31. Type: Native Collector s.n. 1892 “Tringganu’
(Trengganu) (holo K ex SING’)
Synonym nova: Begonia peninsulae Irnscher ssp. peninsulae, Mitt. Inst. Allg. Bot.
Hamburg. 8 (1929) 98. Type: ‘Tringganu’ without collector, number or date (K ex
SING’).
Subspecies tambelanense
Irmscher described this subspecies (a begonia with peltate leaves) from a collection
by Ridley made in 1895 from a plant that was cultivated in the Botanic Garden
Singapore. This, in turn, was originally collected by Pereira from the Tambelan Islands,
NW of Pontianak, Indonesian Borneo.
It is very different from the specimen of his subsp. peninsulae (=B. rajah)
from Trengganu as not only are the leaves peltate with a long acumen but the petiole
and lower surface of the veins are conspicuously hairy. (Those of B. rajah are deeply
cordate with a short acumen and are more or less glabrous). Again, Irmscher included
plants of unknown provenance collected from both Gardens — Nur Begonia No. 2
(SING), Nur Begonia No. 3 (K, SING) both dated March 1917 from the Botanic
‘When Irmscher monographed the begonias of Peninsular Malaysia he borrowed the
entire collection of the Singapore Botanic Gardens (SING) and these he annotated
and cited in his monograph. He did not consult the collection at Kew (K) and none of
their specimens is cited by him. A considerable number of unicate types are no longer
to be found at SING but are now at K. We have no record of how they came to be
transferred but they are definitely the Singapore types as they bear Irmscher’s original
annotations. These are cited as K ex SING in this account.
Begonia peninsulae 63
Gardens Singapore; and Burkill Begonia No. 1 (SING) July 1917, Burkill 6115 (SING)
31 Aug 1920 from the Penang Botanic Garden. These are among a gathering of
begonias by Burkill that included ornamental begonias made over 20 years after the
original Pereira specimen was introduced into Singapore. It is unlikely that the
Tambelan plants had survived so long.
These garden plants differ from the original collection by Ridley in their
more densely pilose petioles and distinctly toothed leaf margin. Irmscher had already
raised the possibility that these were pot plants (Topfpflanzen) and that they differed
from the Tambelan plant in leaf size (6.5—17 x 8.5—24 cm), margin and acumen length.
The Tambelan plant is therefore regarded as a distinct species and is here raised to
specific rank.
To add confusion, Irmscher also recognised a hybrid between the two
subspecies, Burkill 6113 31 Aug 1920 (SING), because he considered it was
intermediate between the two subspecies. This specimen is less hairy and has a less
conspicuously toothed margin than Burkill 6/15 but it does not resemble B. goegoensis
(Burkill 6114) at all.
Begonia tambelanense (Irmsch.) Kiew, stat. nov.
Basinym: Begonia peninsulae Irmsch. ssp. tambelanense Irmsch. Mitt. Inst. Allg. Bot.
Hamburg. 8 (1929) 100. Type: Ridley s.n. 31 Aug 1895 Botanic Gardens Singapore
originally collected by Pereira from Tambelan Is. (holo K ex SING’).
Reference
Irmscher, E. 1929. Die Begoniaceen der Malaiischen Halbinsel. Mitteilung aus dem
Institut fur allgemeine Botanik in Hamburg. 8: 86—160.
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Garden’s Bulletin Singapore 55 (2003) 65—68&
A New Species and a New Combination in Bornean Kopsia
(Apocynaceae: Apocynoideae)
DAVID J. MIDDLETON
Harvard University Herbaria, 22 Divinity Avenue, Cambridge, MA 02138, U.S.A.
Abstract
A new species, Kopsia rajangensis D.J.Middleton, is described and a new combination, Kopsia
pauciflora Hook.f. var. mitrephora (Sleesen) D.J.Middleton, is made.
Introduction
As preparation for an account of the Apocynaceae for the Tree Flora of Sabah and
Sarawak, it is necessary to describe a new species of Kopsia and reduce Kopsia
mitrephora Sleesen to a variety of Kopsia pauciflora Hook.f.
Kopsia rajangensis D.J.Middleton, sp. nov.
Frutex 1.2-4.6 m altus, ramis glabris. Folia elliptico-oblonga, 13.4—32 x 3.8—9.6 cm,
nervis 9—25 paribus. Corollae tubus 21.5—25 cm, lobis 11-14 x 2.7-4.7 mm. Typus:
Borneo, Sarawak, Kapit, Upper Rejang River, Clemens & Clemens 21221 (holo MO;
iso A, BM, BO, NY, SAR)
Small tree or shrub, 1.2—-4.6 m tall. Branchlets glabrous, not or sparsely lenticellate,
terete. Leaves: petiole 4-9 mm long, glabrous; blade papery to subcoriaceous, elliptic
or oblong, apex caudate, base obtuse to cuneate, 13.4—32 x 3.8—9.6 cm, 2.64.3 times
as long as wide, midrib shallowly sunken or raised and with a central groove above,
secondary veins 9—25 with 6—25 mm spacing, 40—60° from midrib, prominent or flat
above, prominent beneath, clearly discernable from tertiary venation above and
beneath, straight or slightly ascending near margin, tertiary venation prominent above,
prominent or flat beneath, reticulate or sub-perpendicular to midrib and oblique to
secondary veins, intramarginal vein looped and inset from margin, glabrous above
and beneath. Inflorescences dichasial and then with cincinnate branches, 4-15 cm
long with axes 14-140 x 1.4—2.1 mm, glabrous to densely puberulent; peduncle 2—76
66 Gard. Bull. Singapore 55 (2003)
x 1.7—3.7 mm; pedicel c. 4 mm long; bracts persistent in inflorescence. Sepals ovate,
apex rounded, 1.5—1.7 x 1.1—1.4 mm, 1.2—1.4 times as long as wide, ciliate, otherwise
glabrous outside and inside. Corolla white; tube 21.5—25 x c. 2.3 mm, pubescent
around stamens and slightly beneath inside, throat pubescent, glabrous outside, 1.8—2 times
as long as lobes, 12.6—16.7 times as long as calyx; lobes elliptic or oblong, apex
obtuse or acute, 11—14 x 2.7-4.7 mm wide, 34.5 times as long as wide, not ciliate,
glabrous outside and inside. Stamens 17-21 mm from corolla base which is 0.79-0.84
of corolla tube length; anthers 1.7—2 x 0.6—0.8 mm, 2.5—2.8 times as long as wide,
1.1—2 mm from corolla throat; filaments c. 0.7 mm long. Ovary 0.9 mm high, glabrous
to densely pubescent; style 20 mm long; pistil head 0.8 mm long. Fruits falcate with
small blunt projection near the base, 15—16 x 4 x 6.5—7 mm; projection 34 mm long.
Distribution: Borneo - SARAWAK: endemic to the Rajang River area.
Habitat: In hill forest and disturbed primary forest on rich clay soil.
Notes: Merrill (1921) compared a specimen of this species, Haviland 3042, to Kopsia
macrophylla Hook.f. but offered no further identification. Timmerman- Van der Sleesen
(1959) was the first to note the peculiarity of the material and compared it to Kopsia
lapidelecta Sleesen and Kopsia tenuis Leenh. & van Steenis, again without making a
recommendation as to what it might be or describing a new species. Anderson (1980)
and Ashton (1988) called this species Kopsia larutensis King & Gamble, a species
known only from Peninsular Malaysia. Kopsia rajangensis differs from K. larutensis
in its larger inflorescences, much larger flowers and relatively wider corolla lobes.
Markgraf (1972) cited the specimens under Kopsia arborea Blume. As can be seen,
there has been remarkably little agreement on the identification of the species around
Kapit on the Rajang River, which the Clemenses spelt ‘Rejang’. Part of the problem
has been the lack of good material.
This new species is probably related to Kopsia lapidelecta and Kopsia
sleeseniana Markgr. It differs from the former in its secondary venation, intramarginal
vein and larger flowers, and from the latter in its smaller flowers and also its venation.
Specimens examined: Sarawak: Kapit, Upper Rajang River, Clemens & Clemens
21211 (SAR), Clemens & Clemens 21221 (A, BM, BO, MO, NY, SAR); Pelagus
Rapids, Ashton $17797 (L); Rajang River, Haviland 3042 (BM, SAR, SING);
Bukit Raya, Smith S27738 (SAR).
Kopsia pauciflora Hook.f. var. mitrephora (Sleesen) D.J.Middleton, comb. nov.
Basionym: Kopsia mitrephora Sleesen, Blumea 10 (1960) 136. Type: G.H.S. Wood
SAN 16118. Borneo, Sabah, Lahad Datu, Path between Sungai Sabahan and Sungai
Bornean Kopsia 67
Dok (holo L; iso BRI).
Notes: Kopsia mitrephora was formally described by Timmerman-Van der Sleesen
(1960) following on from her earlier, but not validly published, account of the species
(1959). In the earlier work she distinguished K. mitrephora on the basis of its stamens
inserted in the lower part of the corolla tube, a character unusual in the genus, and on
its delicate leaves. However, with the many more specimens available now, it is clear
that the vegetative characters in this species are extremely variable and closely parallel
the vegetative variation found in K. pauciflora. These variations include the size of
the wings or angles on the stems, the thickness of the leaves, the length of the petiole
and the degree of prominence of the venation. Despite long attempts to put some sort
of order to this variation, and indeed there are character states concentrated in some
geographical areas, no further taxa could be distinguished in either species. In addition,
it became increasingly clear that the only character to distinguish K. pauciflora and
K. mitrephora was the site of insertion of the stamens and that otherwise both species
were remarkably similar, including in fruit characters. Therefore, varietal status is
appropriate for Kopsia pauciflora var. mitrephora.
Acknowledgments
The study was partly carried out under the sponsorship of The Tree Flora of Sabah
and Sarawak Project and funded by the Ministry of Science, Technology and
Environment of Malaysia through IRPA Funding No. 30—08—01—01-005. The curators
of the herbaria at BM, BO, L MO, NY, SAR and SING are thanked for either a loan of
material or for their hospitality whilst the material was consulted.
References
Anderson, J.A.R. 1980. A Checklist of the Trees of Sarawak. Forest Dept. Sarawak,
Malaysia.
Ashton, P.S. 1988. Manual of the Non—dipterocarp Trees of Sarawak. Forest Dept.
Sarawak, Malaysia.
Markgraf, F. 1972. Florae Malesianae Praecursores LIII. Apocynaceae II. 6. Urnularia,
7. Willughbeia, 8. Kopsia. Blumea. 20: 407-425
Merrill, E.D. 1921. A Bibliographic Enumeration of Bornean Plants. Fraser & Neave,
Singapore.
Timmerman-Van der Sleesen, E.H.L. 1959. Preliminary revision of the Malaysian
species of the genus Kopsia (Apocynaceae). Flora Malesiana Miscellaneous
Records. 1: 1-15
68 Gard. Bull. Singapore 55 (2003)
Timmerman-Van der Sleesen, E.H.L. 1960. In C.G.G.J van Steenis, Miscellaneous
Botanical Notes X. Blumea. 10: 136—139
Garden’s Bulletin Singapore 55 (2003) 69-72
New Species and Varieties of Symplocos (Symplocaceae)
from Borneo
K.G. PEARCE
32, Lorong Kumpang 4, Kuching, Sarawak, Malaysia
Abstract
A new species of Symplocos, S. buxifolioides K.G.Pearce and two varieties, S. iliaspaiensis
Noot. var. pedunculata K.G.Pearce and S. laeteviridis Stapf var. alabensis K.G.Pearce are
described from Borneo.
Introduction
In his account of the Symplocaceae for the Flora Malesiana region, Nooteboom (1975,
1977, 1984, 1986, 1989) recognised 30 species with 19 varieties of the genus Symplocos
Jacq. in Borneo. In the process of revising and preparing the manuscript for the Tree
Flora of Sabah and Sarawak, one new species, Symplocos buxifolioides, and two
varieties, S. iliaspaiensis var. pedunculata and S. laeteviridis var. alabensis are
recognised. Descriptions and brief notes on the geographical distribution and
morphological characters distinguishing these new taxa from those of previously
known are given.
1. Symplocos buxifolioides K.G.Pearce, sp. nov.
Symplocos buxifoliae Stapf similis a qua differt ramulis distincte modo gallice et
anglice ‘zigzag’ et margine foliae revoluto praeter interdum ad apicem et e basi foliorum
non altae obscure denticulatae. Typus: Chew & Corner RSNB 5882, Borneo, Sabah,
Mt. Kinabalu, Mesilau (holo SING; iso SAN).
Tree to 10 m tall, 10 cm diam. Twigs weakly, irregularly curved, smaller ones distinctly
zig—zag with nodes closely set, slender, up to 4 mm diam., dark brown to black; young
parts longitudinally ridged and sparsely appressed long—pilose; older parts becoming
horizontally cracked, glabrescent, without prominently raised leaf scars. Leaves
spirally arranged, coriaceous, glabrous above, hairy to virtually glabrous below, drying
olive—brown above, yellow—green below; blade broadly ovate—elliptic, 2.54.5 x 1-3
70 Gard. Bull. Singapore 55 (2003)
cm, base shortly attenuate, margin revolute except sometimes at apex, shallowly,
obscurely toothed from near base, apex acute, shortly apiculate; midrib narrowly
channelled above; /ateral veins (6—)7-& pairs, joining to form an intramarginal vein;
intercostal venation reticulate, raised above and prominent below; petiole 3.5—5 mm
long, sharply flanged to base. Inflorescences racemous, borne in the axils of upper
leaves and below on leafless twigs; axis 2—5(—9) mm long, shortly, moderately dense
appressed—pilose; bracts early caducous. Flowers 1-3; pedicels to 0.5 mm long;
calyx tube c. 0.3 mm long, lobes broadly ovate, chartaceous, ciliate almost to apex,
moderately dense to sparsely appressed—hairy, c. 1 mm long; corolla white, glabrous,
3.5—4 mm long; ovary sparsely to densely appressed hairy, 1.5 mm high. Fruits
solitary on a pedicel to 9 mm long, ovoid to curved ellipsoid, 1-1.5 x 0.5—0.7 cm,
hardly to shallowly ridged, sparsely appressed—pilose to glabrescent, drying yellowish
brown, with persistent and erect calyx lobes.
Distribution: Endemic in Borneo. Known only from Mt. Kinabalu, Sabah.
Ecology: Upper montane forest at 2400-3660 m altitude.
Other specimens examined: SABAH. Mt. Kinabalu: Hotta 3840 (SAN), Chew &
Corner RSNB 5911 (SAN), Mikil & Aban SAN 46593 (SAN) and Aban et al. SAN
54254 (SAN).
Notes: Symplocos buxifolioides is similar to S. buxifolia Stapf but differs from the
latter in its slender twigs, which are distinctly and shortly zig—zag between the closely
set nodes; young parts of the twigs sparsely appressed pilose; broadly ovate—elliptic
leaves (not obovate—elliptic as in S. buxifolia) with revolute and closely, obscurely
denticulate margin from near the base, acute to shortly apiculate apex (not obtuse to
rounded), and 6—8 pairs of lateral veins (not 4—6); inflorescences borne on leafless
twigs as well as in the axils of upper leaves; calyx tube c. 0.3 mm long, ciliate almost
to the apex (not sometimes ciliate at base), and moderately densely to sparsely
appressed—hairy (not glabrous); and sparsely to densely appressed—hairy ovary (not
glabrous). This species is named for its close morphological similarity to S. buxifolia.
2. Symplocos iliaspaiensis Noot. var. pedunculata K.G.Pearce, var. nov.
A var. iliapaiense differt racemis base ramosis ad 6.5 cm longis floribus (fere)
sessilibus ad 2 mm pedicellatis. Typus: Fedilis SAN 95642, Borneo, Sabah, Kalabakan
district (holo SAN; iso K, KEP, L, SAR, SING).
Tree to 13 m tall. Leaves 7.5—20 x 2.5—6 cm, base cuneate, apex with acumen to 1.4
cm long. Racemes basally branched, to 6.5 cm long. Flowers virtually sessile or
Symplocos from Borneo al
with short pedicel to 2 mm long.
Distribution: Endemic in Borneo. Sabah (Kalabakan, Sandakan and Tawau districts)
and Brunei Darussalam.
Ecology: In primary and logged—over mixed dipterocarp forest at altitudes to 80 m,
on flat or undulating lands and ridges.
Other specimens examined: SABAH. Sandakan district, Sigin SAN 56798 (KEP, SAR,
SING), Leopold & Kodoh SAN 81396 (SAN, SAR), Leopold & Kodoh SAN 81424
(SAN, SAR); Lahad Datu Sinanggul SAN 56991 (SAN, SAR); Kinabatangan Meijer
SAN 136314 (SAN), Pensiangan district Goh SAN 141166 (KEP, SAN). BRUNEI:
Belait Wong WKM 1079 (SAN).
Notes: This new variety may reach 13 m tall (not 3.3 m tall as in the typical variety).
It is vegetatively very similar to var. iliaspaiensis but can be distinguished by its
basally branched raceme or unbranched spike or fascicle of racemes 1.5 to 6.5 cm
long bearing virtually sessile to shortly pedicellate (to 2 mm long) flowers. In contrast,
var. iliaspaiensis has fascicles of sessile flowers with the axis entirely absent. This
variety is named for its pedunculate inflorescences.
3. Symplocos laeteviridis Stapf var. alabensis K.G.Pearce, var. nov.
Symplocos laeteviridis Stapf var. mjobergii (Merr.) Noot. similis praeter ramunculos
graciliores cum paginis infernis foliorum omnino glabros, foliis olivaceis, parvioribus,
lanceolatis, ad 6.3 cm longis et 2.2 cm latis. Typus: Madani & Majawat SAN 119252.
Borneo, Sabah, Penampang district, Gunung Alab (holo SAN, iso KEP).
Glabrous treelet to 2 m tall. Twigs slender, glabrous. Leaves drying olive green;
blade glabrous below, lanceolate, to 6.3 x 2.2 cm, base cordate; lateral veins 10 pairs;
petiole c. 1 mm long.
Distribution: Endemic in Borneo. Sabah, known only from Gunung Alab (the type
specimen and Kamaruddin KMS 1431 (SAN, UKMS).
Notes: The new variety is similar to Symplocos laeteviridis Stapf var. mjobergii (Merr.)
Noot. except that the twigs and lower leaf surface are entirely glabrous and it has
more slender twigs and smaller (to 6.3 x 2.2 cm) lanceolate leaves that dry olive
green. It is named for the locality where it was collected.
7? Gard. Bull. Singapore 55 (2003)
Acknowledgements
The work was carried out under the Tree Flora of Sabah and Sarawak Project and
funded by IRPA Research Grant 30—08—01—01—005. The curators of the herbaria at
K, KEP, SAN, SAR and SING are thanked for permission to examine specimens in
their care as is H.P. Nooteboom for translating the diagnoses into Latin.
References
Nooteboom, H.P. 1975. Revision of the Symplocaceae of the Old World, New
Caledonia excepted. Leiden Botanical Series. 1: 33-335.
Nooteboom, H.P. 1977. Symplocaceae. Flora Malesiana. I, 8: 205-274.
Nooteboom, H.P. 1984. Symplocos (Symplocaceae) from Bukit Raya. Blumea. 30:73-74.
Nooteboom, H.P. 1986. Additions to Bornean Symplocaceae. Blumea. 31: 277-279.
Nooteboom, H.P. 1989. Addenda, corrigenda et emendanda. Flora Malesiana. 1,10: 719.
Garden’s Bulletin Singapore 55 (2003 ) 73-88
Five New Begonia species (Begoniaceae) from the Niah
National Park, Sarawak, Malaysia
KATHARINE G. PEARCE
DANIDA/SWMPI Project, Sarawak Forest Department, Kuching, Malaysia
Abstract
Five new Begonia species, Begonia kachak K.G.Pearce, B. kasutensis K.G.Pearce, B. niahensis
K.G.Pearce, B. stichochaete K.G. Pearce and B. subisensis K.G.Pearce, were collected from
limestone habitats in the Niah National Park. A key, descriptions and illustrations are provided
Introduction
Sarawak’s varied geology includes a number of isolated limestone outcrops dating
from a range of geological epochs and varying in size. In north Sarawak the large
Melinau formation falls partly within the Gunung Mulu National Park, while in west
Sarawak there are a number of limestone outcrops around Bau and Serian and south
towards Tebedu. Other exposures are scattered 1n the interior of Sarawak, for example,
at Bukit Sarang (Tatau), Ulu Kakus and in the Middle Baram. Between Bintulu and
Miri, in the Niah area about 16 km from the coast, there is an isolated limestone
massif known as the Subis massif. In 1974, this massif and surrounding forest was
constituted as the Niah National Park.
The Bau limestone outcrops, being relatively accessible from Kuching, have
been a favourite destination for plant collectors since the mid 19" Century and the
Mulu limestone was the focus of intensive botanical collection during the Sarawak
Forest Department and Royal Geographic Society Expedition in 1978. In contrast,
the Subis massif was, until recently, somewhat neglected by plant collectors. G.D.
Haviland, Curator of the Sarawak Museum, Kuching, and C. Hose, District Officer
in Baram and Sibu, together made the first plant collections there in 1894. In 1932,
P.M. Synge, a member of the 1932—-33 Oxford University Expedition to Sarawak,
visited Subis, while in 1954, W.M.A. Brooke and Ahmad independently made
collections in the area under the Sarawak Museum label. In 1962, B.L. Burtt with
P.J.B. Woods and Chew W.L. collected there as did H.P. Fuchs in 1963 and G.
74 Gard. Bull. Singapore 55 (2003)
Alphonso and Samsuri Ahmad of the Singapore Botanic Gardens in 1965. The Botany
Unit staff, Sarawak Forest Department, made collecting trips to Niah on a number of
occasions from the early sixties onwards, including J.A.R. Anderson, who had a
particular interest in the limestone flora of Sarawak (Anderson, 1965) and who
collected there in 1966 and 1972. Their collections included specimens of the five
begonia species described below but which until now remained unnamed.
All five species are endemic to the Subis limestone.
The highest point of the Subis limestone massif is G. Subis. Bukit Kasut and
Gua Pangomah are located on the west of the massif. G. Brangin and G. Bekajang are
two discrete hills to the north and northeast of the massif, respectively. The latter is
the site of the Niah Great Cave and the Painted Cave, which are important
archaeological sites.
The botanical field investigations in Niah National Park were carried out as
part of the Sarawak Forest Department/DANIDA Project ‘Support to Wild Life Master
Plan Implementation through the Improved Management of Totally Protected Areas
in Sarawak, Malaysia’ and enabled the five Begonia species to be re-collected from
the limestone habitats and for detailed observation to be made.
Key to Begonias in the Niah National Park
la, Leat- ovate, oblong-ovate of Obovate = oo... 22.6 seaeese cena teen eee
ib.) Leafwzeniform:to:sub+orbicular, sejcnic. 4 nell \enlineg beta eee
2a... ~Leaf 1.5 times.or moreastlong:as broad auiiiwe elim lab nehenel cee
2b. Leaf less than 1.5 times as long as broad ...................5..2.....+. 2. B. niahensis
3a. Leaf with lines of stiff hairs between veins ...................... 3. B. stichochaete
3b. Leaf without lines of stiff hairs between veins ...................... 5. B. subisensis
4a. Leaf more than 7 cm long and wide .....d28. Jena! dole edi.od bee
4b. Leaf less than 7 cm long and wide ...............ssccsseeeeeeee 4. B. Kasutensis
I. Begonia kachak K.G.Pearce, sp. nov.
Section Petermannia
A Begonia speluncae Ridl. foliis majoribus ob internodia longiora non in fasciculis
dispositis nec peltatis nec glabris differt. Typus: Great Cave, Niah, Miri District /.A.R.
Anderson, S. Tan & E.Wright S26074 (holo SAR).
Begonias from Niah National Park WS
Figure 1. Begonia kachak
A Habit, B Detail of leaf margin, C Male bud, D Male flower, E Androecium, F Stamens, G
Female flower, H Style and stigma. (from $26074).
76 Gard. Bull. Singapore 55 (2003)
Figure 1
Creeping herb or low root climber, roots fibrous, produced at the nodes in contact
with the substrate. Stem and petiole with minute appressed to oblique brown hairs;
distal end of young petiole, main veins on abaxial surface, margins of young leaf and
main vein of stipule with stiff pink-brown trichomes to | mm long. Leafy stem 6-28 cm
long, 3 mm diam., little branched, internodes 1—5 cm long. Stipules broadly triangular
with a distinct midrib, 4.5 x 7 mm, margin entire, prolonged into an acuminate apex,
persistent. Leaves alternate; petiole 1.8 cm in upper leaves, to 17 cm in lower leaves;
lamina minutely papillose above and below, upper surface somewhat lustrous, olive-
green to yellowish green especially near margins, lower surface pale green or in some
specimens brownish pink with pale green veins, oblique, reniform to sub-orbicular,
7.5—12.5 x 7.5—15 cm, base cordate with the basal lobes hardly overlapping, rounded,
0.24.7 cm long, margin irregularly and indistinctly serrate, teeth setose, apex acute
to acuminate, venation pinnate-palmate, main veins 4 pairs, branching about half
way to the margin, with 2 more veins in the broader and one in the narrower basal
lobe, sunken above, prominent beneath. /nflorescences axillary, protogynous, obliquely
erect cymose panicles, 8.5—29 cm long, of which peduncle is up to 14 cm long, with
one or two female flowers proximally and many male flowers distally; bracts pink,
similar to stipules, 6.5 x 3 mm, persistent, bracteoles pale pink-brown, broadly ovate,
4 x 3 mm, apex setose, persistent. Male flowers with pink pedicel to 7.5 mm long,
tepals 2, white translucent margins pink before anthesis, minutely papillose, orbicular,
7x 5.5 mm, margin entire, apex rounded, stamens yellow, c. 29, in a spherical cluster,
joined in a torus c. 3 mm diam., filaments to 1 mm long, anthers pale yellow, obovate,
c. 0.7 mm long, emarginate. Female flowers on pedicels 5.5 mm long, minutely hairy;
ovary pink, oblong-obovoid, 10.5 x 7 mm, wings pink, 3, subequal, c. 1.5 mm wide;
tepals 5, white translucent or palest pink, with scattered minute, deeper pink hairs,
unequal, oblong with rounded apex, to 9.5 x 5.5 mm, margin entire; styles 3, free,
each 2 mm to bifurcation from where papillose stigmatic surface forms a continuous
twisted band. Fruits with decurved pedicel, 1—1.9 cm long; capsule asymmetric,
broadly oblong-elliptic, to 1.9 cm x 1.6 cm, minutely hairy, papillose, locules 3, each
with 2 placentae, dehiscing between wing and locule, wings 3, unequal, thin, narrowed
to base and apex, two narrower wings to 5 mm wide, broader wing to 7 mm wide,
style caducous. Seeds cylindrical, c. 0.3 x 0.2 mm, surface areolate, areoles more or
less equal sided at base and apex and elongate along seed axis.
Distribution: Borneo — SARAWAK: endemic in the Subis limestone (G. Brangin,
Niah Great Cave and Gua Pangomah).
Habitat: Locally abundant in the shelter of overhangs in somewhat dry areas or
Begonias from Niah National Park Vl
creeping up rocks at cave mouths, infrequently at level ground in deep shade adjacent
to limestone outcrops.
Notes: This is a conspicuous begonia with large shiny leaves, commonly bearing a
cluster of delicate, pale pink male flowers. It is easily observed on limestone outcrops
along the Plank Walk to the Niah Great Cave and around the cave mouth. It takes its
name from kachak, the Malay word meaning handsome.
Specimens examined: SARAWAK: Gunung Subis, Niah B.L. Burtt & P.J.B. Woods
B2010 (SAR), Mohidin S2 1603 (SAR); Great Cave K.G. Pearce, Bibian Diway, Saupel
Atot & Dami Jude $78536 (SAR), Jemree Sabli S89062 (SAR); G. Brangin, Ulu Sg
Subis Yii Puan Ching S40168 (SAR), Gua Pangomah J.A.R. Anderson S31691 (SAR,
SING), Near Sg Subis K.G. Pearce $89463 (SAR); Niah Caves Alphonso & Samsuri
A217 (SING), A222 (SING), A248 (SING).
2. Begonia niahensis K.G.Pearce, sp. nov.
Section Petermannia
A Begonia congesta Ridl. habitu erecto vel effuso (non cauli longo), capsula oblongo-
ovoidea (non longa oblongaque) differt. Typus: Niah Caves B.L. Burtt & P.J.B. Woods
B 2009 (holo SAR).
Figure 2
Herb with stems to 75 cm long, branching near base with some spreading horizontally,
some vertical and some drooping and then curving upward. Young stems and petioles
with minute, erect translucent hairs. Older stems brown and semi-woody, to 1 cm
diam., young stems brownish pink, internodes 2—8.5 cm long. Stipules green flushed
brownish pink at centre, plicate, broadly ovate, 14 x 7 mm, margin entire with
translucent hairs at base, apex acute, persistent. Leaves spirally arranged; petiole from
c. 5 cm long in the upper leaves to 22 cm in the lower leaves, deep brownish pink;
lamina medium green above, paler below, with veins on both surfaces deep pink at
base and at the point where they branch, pinkish between, minutely papillose with a
velvety appearance above, translucent hairy beneath, oblique, broadly ovate, 15—20 x
12—16.6 cm, base cordate with basal lobes not or hardly overlapping, rounded, to 9
cm long, margin irregularly and indistinctly serrate, translucent hairy, teeth with
translucent setae, apex acute, venation pinnate-palmate, main veins 4—5 pairs,
branching near base, with 2 or 3 smaller veins in broader basal lobe and 1 or 2 in
narrower basal lobe. /nflorescences axillary, protogynous, obliquely upright, cymose
panicles 3—8 cm long, of which the peduncle is 2.5—-6 cm long, with | or 2 pairs of
female flowers at the base with many male flowers tightly congested distally, bracts
78 Gard. Bull. Singapore 55 (2003)
plicate, pale green, broadly ovate, to 2—2.3 x 1.2 cm, persistent. Male flowers opening
one at a time, pedicel obscured by congested bracts and male flower buds, tepals 2,
white translucent, margins entire, orbicular, c. 6 x 7 mm, apex rounded; stamens
yellow > 30, in hemispherical cluster, joined in a torus c. 1 mm diam., filaments c. 0.5 mm
long, anthers yellow, emarginate, 0.8 mm long, opening by apical slits. Female flowers
with pedicel c. 9 mm long; ovary yellow-green to pink, obovoid, 19 x 7 mm, wings 3,
pale green, subequal, 1.5 mm wide; tepals 5, white translucent, subequal, broadly
elliptic-obovate, 13 x 7 mm, margins entire except at fringed, obtuse apex, outer
surface and margins minutely translucent hairy; styles 3, c. 1 mm long; stigmas
bifurcated, stigmatic surface a minutely papillose, orange yellow spiral band. Fruits
with a stiff, decurved pedicel, c. 1.5 cm long, pale green flushed pink at base; capsule
oblong, to 3.2 x 1.4 cm, sparsely translucent-hairy, locules 3, each with 2 placentae,
dehiscing between wing and locule, wings 3, subequal, thin, broadest at apex, c. 4 mm
wide, style caducous. Seeds cylindrical to obovoid, c. 0.3 x 0.2 mm, surface areolate,
areoles more or less equal sided at base and apex and elongate along seed axis.
Distribution: Borneo - SARAWAK: endemic in the Subis limestone (G. Brangin,
Niah Great Cave and Painted Cave areas).
Habitat: Below cave mouths, rooted in pockets of soil on ledges and in crevices on
steeply sloping limestone rock or in light shade under canopy of small trees.
Notes: This robust and handsome begonia has large, asymmetric, velvety hairy leaves
that sometimes have a bluish cast. The leaves are positioned more or less vertically
against the limestone rock face in somewhat sheltered sites. It can be observed on
limestone outcrops along the Plank Walk to the Painted Cave. It has been named for
the type location - the Niah Caves. Its habit of drooping and upcurving stems and
handsome, velvety leaves held vertically, give this species potential as an ornamental
plant.
Specimens examined: SARAWAK: G. Brangin Yii Puan Ching $40166 (SAR); Great
Cave J.A.R. Anderson, S. Tan & E. Wright S26075 (SAR, SING), K.G. Pearce, Bibian
Diway, Saupel Atot & Dami Jude §78537 (SAR); Near Sg Subis K.G. Pearce S89460
(SAR).
3. Begonia stichochaete K.G.Pearce, sp. nov.
Section Petermannia
A Begonia congesta Ridl. foliis angustioribus seriebus setarum inter venas gerentibus,
capsulis longioribus oblongis (non oblongo-ovoideis) differt. Typus: Subis, Niah Caves
Ahmad 14 (holo SAR; iso SING).
Begonias from Niah National Park
Figure 2. Begonia niahensis
A Habit, B Detail of leaf undersurface, C Detail of leaf margin and upper surface, D Male
bud, E Male flower, F Androecium, G Stamens, H Female flower, I Style and stigma from
above, J Style and stigma. (A—C from Burtt 2009, D-I from $89460).
79
80 Gard. Bull. Singapore 55 (2003)
Figure 3
Erect herb rooting at the base of the stem and at nodes along the horizontal rhizome
which creeps below the soil surface and gives rise to shoots at intervals. Indumentum
of stiff trichomes to 2.2 mm long, each on a raised conical base, pale brown to pinkish
brown to deep brownish red on stem, young petiole and veins on lower and upper
lamina surfaces, dense on young stem and petiole, less so on stipule margin and
sparse on outer surface of stipule midrib and on the main veins of the upper lamina
surface, in a row or rows between each pair of veins and on and between the teeth of
the dentate margin, moderately dense on the main veins of the lower leaf surface.
Upright stem to c. 42 cm long, 5—9 mm diam., unbranched, internodes 2.5—5 cm long
to 8 cm long at base of stem. Stipules oblong-ovate with a distinct midrib, 17 x 11 mm,
margin entire, apex setose, caducous. Leaves spirally arranged; petiole 0.7 cm in
upper leaves, to 2.5 cm in lower leaves; lamina at first yellowish green then medium
green on upper surface (rarely dark blackish-green), lower surface paler yellowish
green or olive-green or deep pink or red, asymmetric, irregularly obovate, 11-19 cm
long, narrower side to 2.5 cm wide towards apex, curving inwards towards base and
hardly or not lobed at base, broader side 6—7.5 cm wide with a cordate base and a
rounded basal lobe to 17 mm long, margin irregularly serrate (appearing almost
praemorse), each tooth setose, with several setae between each pair of teeth, apex
toothed, venation pinnate, c. 4 pairs of veins, branching with another 1-3 veins in
basal lobes, impressed above, prominent beneath. /nflorescences axillary, protogynous,
erect cymose panicles to 3.4 cm long, of which the peduncle is 2.3 cm, with 1—2
female flowers at base with distally many male flowers crowded in a terminal cluster;
bracts similar to stipules, deep red, 13 x 10 mm, persistent; bracteoles pale brown to
deep pink, broadly ovate, to 12 x 12 mm, apex setose, persistent. Male flowers witha
green to deep pink pedicel with deep pink bristles, 7 mm long; tepals 2 or 4, the outer
pair translucent white flushed pink at margin and base, the inner pair translucent
white, at first finely hairy on outer surface near base, broadly elliptic, c. 4 x 3 mm,
margin entire, apex rounded; stamens pale yellow, c. 32, in an ovoid cluster, joined in
a torus c. | mm diam., filaments to | mm long, anthers obovate, to 1.5 mm long,
shortly apiculate, opening by terminal pores which develop into slits. Female flowers
solitary or in pairs; pedicels 5 mm long, medium pink with minute appressed to slightly
oblique hairs; ovary pale pink tinged green with deep pink bristles, oblong-obovoid,
to 24 x 16 mm, wings 3, deep pink to translucent white with margins flushed pale
pink, subequal, c. 7 mm wide; tepals 5, translucent white flushed pale pink at margins
or palest pink, deeper pink at base and on marginal teeth with scattered deeper pink
hairs, on outer surface sparsely hispid, oblong with rounded apex, inner two larger, c.
6 x 2-4 mm, margin toothed from near base to fringed apex; styles 3, free to base, 3 mm
long, bifurcating; stigmas forming a continuous twisted band, yellowish-green to bright
yellow. Fruits with stiff, decurved pedicels, 5—9 mm long; capsule oblong,to 29 x 18 mm,
81
Begonias from Niah National Park
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82 Gard. Bull. Singapore 55 (2003)
sparsely hispid, minutely papillose, locules 3, each with 2 placentae, dehiscing between
wing and locule, wings 3, subequal, thin, narrowed to base, truncate distally, two
narrower wings to 5 mm wide at midpoint and 6 mm at apex, broader wing to 6 mm
wide at midpoint to 7 mm at apex, style caducous. Seeds cylindrical-spherical, c. 0.3
x 0.25 mm, surface areolate, areoles more or less equal sided at base and apex and
elongate along seed axis.
Distribution: Borneo —-SARAWAK: endemic in the Niah National Park (Kuala Subis,
the Sekaloh area and near the trail to Bukit Kasut).
Habitat: This species occurs both in limestone habitats and undisturbed Mixed
Dipterocarp Forest on loamy soil or brown clay as well as in secondary forest on
alluvial soil. On limestone, it grows on low outcrops with little soil and in soil at the
base of limestone cliffs in deep shade and damp conditions, where it is locally frequent.
Notes: This erect begonia is characterised by its bristly leaves bearing rows of stiff
hairs between the veins. It grows in shady sites, both on or near limestone outcrops
and in forest away from the limestone. The species shows striking variation with the
upper leaf surface ranging from mid-green to dark blackish-green. It is named for its
bristly leaves.
Specimens examined: SARAWAK: Niah National Park Bernard Lee S40075 (SAR),
Kuala Niah near confluence with Sg Subis K.G. Pearce, Bibian Diway, Saupel Atot &
Dami Jude S78539 (SAR); Route to Bukit Kasut K.G. Pearce & Narawi b. Johari
S75597 (SAR); Sekaloh Jemree Sabli S8S9014 (SAR), K.G. Pearce S589267 (SAR).
4. Begonia kasutensis K.G.Pearce, sp. nov.
Section Petermannia
A Begonia conipila Irmsch. ex Kiew trichomatibus brevioribus sine basibus conicalibus,
foliis suborbicularibus (non valde asymmetricis), punctis maculisque praesentibus,
inflorescentiis contra folia insertis differt. Typus: Great Cave, Gunong Subis, Niah,
Miri District J.A.R. Anderson S31940 (holo SAR; iso SING).
Figure 4
Creeping herb with fibrous roots produced at the nodes. Stem and petiole with white
minutely appressed hairs on distal end of petiole; main veins, lamina undersurface
and margin with scattered pale orange-brown hairs to 0.7 mm long, more or less
regularly arranged. Leafy stem toc. 15 cm long, 1.5 mm diam., not or hardly branched,
internodes to 4 cm long. Stipules oblong-ovate with a distinct midrib, to 9 x 4mm,
Begonias from Niah National Park 83
Figure 4. Begonia kasutensis
A Habit, B Detail of leaf margin, C Leaf showing variegation, D Male bud, E Male flower,
F Stamens. (A & B from $31940, C from $27269, D-F from $3/940).
84 Gard. Bull. Singapore 55 (2003)
margin entire, apex setose, caducous. Leaves spirally arranged; petiole 1.5 cm in
upper leaves, to 3.8 cm in lower leaves; lamina minutely papillose above and beneath,
variegated, upper surface slightly lustrous, olive to dark green with irregular spots
and blotches of pale grey-green between veins, lower surface reddish, sometimes
light green towards base, slightly asymmetric, sub-orbicular, 4.7—6.2 x 3.9-5.5 cm,
base cordate with rounded basal lobes not overlapping, to 8 mm long, margin
irregularly, indistinctly and distantly serrulate, apex cuspidate, venation palmate, main
veins 2 pairs, the two ‘middle’ ones branching near mid-point or towards margin,
with 1—2 veins in each basal lobe, sunken above, slightly prominent beneath.
Inflorescences \eaf-opposed, protogynous, racemes of cymules, to 11.5 cm long,
including peduncle to 5.7 cm, with a single female flower at base and many male
flowers distally; bracts similar to stipules, 4 x 2.5 mm, caducous; bracteoles pale
green flushed pink, broadly ovate, 3.5 x 2 mm, apex setose, caducous. Male flowers
with pink pedicel to | cm long; tepals 2, palest pink translucent, suborbicular, c. 7 x 4
mm, margin entire, apex rounded; stamens yellow, c. 20, ina more or less hemispherical
cluster, joined in a torus 0.3 mm diam., filaments to c. 1 mm long, anthers bright
yellow, c. 0.5 mm long, opening by pores, obovate, emarginate. Female flower not
seen. Fruits with stiff, decurved pedicel, 7 mm long; capsule suborbicular, 1.6 x 1.2
cm, locules 3, each with 2 placentae, dehiscing between wing and locule, wings 3,
unequal, thin, narrowed to base, truncate distally, the two narrower wings c. 1.5 mm
wide, broader wing c. 2.5 mm wide, style caducous. Seeds cylindrical, c. 0.3 x 0.25
mm, surface areolate, areoles more or less equal sided at base and apex and elongate
along seed axis.
Distribution: Borneo — SARAWAK: endemic in the Subis limestone (Niah Great
Cave, Bukit Kasut and G. Subis).
Habitat: Infrequent, on limestone rock faces in shaded and dry crevices or in soil
between limestone rocks on slopes of limestone hills up to where the canopy cover
starts to thin out.
Notes: This decorative begonia is characterised by its diminutive, sub-orbicular
variegated leaves that have a pink undersurface, and by its delicate inflorescences. Its
small size and elegant features give it potential as an ornamental species. It has
been named for Bukit Kasut, one of the locations where it occurs.
Specimens examined: SARAWAK: Subis Ahmad No. 3 (SAR, SING); Niah Cave
Ahmad No. 65 (SING); Gunung Subis Jemree Sabli S89049 (SAR); Southern slopes
of G. Subis, near Sekaloh River S. Tan & E. Wright § 27269 (SAR, SING); Route to
Bukit Kasut K.G. Pearce & Narawi b. Johari S78596 (SAR).
Begonias from Niah National Park 85
5. Begonia subisensis K.G.Pearce, sp. nov.
Section Petermannia
A Begonia pendula O. E. Schulz habitu majore caulibus erectis rigidisque, foliis setis
inter dentes setiferos carentibus, fructibus latioribus oblongis differt. Typus: G. Subis
‘in Sekaloh River’, Niah, Miri District /.A.R. Anderson, S. Tan & E. Wright S27574
(holo SAR; iso SING).
Figure 5
Cane-like, semi-herbaceous begonia with woody stems ascending, then held at 45°,
drooping at apex, arising at intervals from a woody, branched rhizome c. 7-9 mm
diam. bearing roots at the nodes, some stem bases bearing adventitious roots. Stem
and petiole with minutely appressed brown hairs. Stems to >76 cm long, from 6 mm
diam. at base to 1.5 mm at apex, bearing leaves only near apex, unbranched or with
few branches, internodes 1.2 to >5 cm long. Stipules lanceolate, without a distinct
midrib, 5.5 x 1.5 mm, margin entire, apex setose, caducous. Leaves spirally arranged;
petiole 1—2.5 cm; lamina minutely papillose above and below, upper surface with a
reflective sheen, medium to olive-green (those of young vegetative shoots variegated,
olive-green with irregular spots of pale grey-green between veins), lower surface dull
green or pinkish between veins to deep pink near veins, asymmetric, oblong-ovate, to
c. 11.3 x 3.64.1 cm, lower half of leaf somewhat inwardly curved, base cordate with
a rounded basal lobe to 1.3 cm long on one side, cuneate or slightly lobed at base on
the other side, margin irregularly serrate, apex acute, venation pinnate-palmate, veins
3-4 pairs, branching except for the outermost two or three, flat or very slightly raised
above, deep pink and prominent beneath. /nflorescences axillary, protogynous, cymose
panicles held obliquely above the subtending leaf, axis pale to deep brownish pink, to
7.5 cm long with | or 2 female flowers at the base and up to 10 male flowers distally
or sometimes, in the lower leaf axils, by reduction, only the female flowers present;
bracts similar to stipules, 4 x 2.5 mm, caducous; bracteoles pale green flushed pink,
lanceolate, 1.5 x 6 mm, apex setose, caducous. Male flowers with pink pedicel to 9
mm long; tepals 2, translucent white to palest pink, some flushed deeper pink at base,
elliptic, 6.5 x 4mm, margin entire, apex rounded; stamens yellow, to 26, in a more or
less obovoid cluster, joined in a 1 mm-diam. torus, filaments c. 1 mm long, anthers
yellow, obovate, emarginate. Female flowers on a peduncle to 3 mm long; pedicels
deep pink to 2.5 cm long, glabrous; ovary oblong, truncate, 1.6 x 2 cm, wings 3,
subequal, c. 6 mm wide; tepals 5, palest pink flushed deeper pink at base, ovate, the
outer two smaller than the inner three, c. 9 x 5 mm, margin entire, apex obtuse, glabrous;
styles 3, bifurcated, 2-3 mm long to bifurcation, stigmatic surface orange with fine
colourless papillae forming a continuous, twisted band. Fruits with flexuous pedicels,
to 3 cm long; capsule oblong, 14 x 23 mm, locules 3, equal, each with 2 placentae,
dehiscing between wing and locule, wings 3, unequal, thin, narrowed to base, truncate
distally, two narrower wings 10 mm wide, broader wing 11.5 mm wide, style caducous.
86 Gard. Bull. Singapore 55 (2003)
Seeds cylindrical, c. 0.4 x 0.25 mm, surface areolate, areoles more or less equal sided
at base and distally, and elongate along seed axis above base.
Distribution: Borneo — SARAWAK: endemic in the Subis limestone (Niah Great
Cave and Bukit Kasut).
Habitat: Locally frequent on limestone rock faces and the steep slopes of limestone
hills in deep litter layer or between limestone rocks where the canopy cover starts to
thin out.
Notes: The leaves of this attractive cane begonia have a reflective sheen on the upper
surface and are deep pink below. The young plant has variegated leaves. The
inflorescence 1s delicate and, in fruit, the capsule dangles on a long, flexuous pedicel.
It has been named for the type locality, G. Subis.
Specimens examined: SARAWAK: Great Cave J.A.R. Anderson $31939, Bukit Kasut
K.G. Pearce, Bibian Diway, Saupel Atot & Dami Jude S78538 (SAR).
Acknowledgements
This paper is based on botanical field investigations at the Niah National Park carried
out under the Sarawak Forest Department/DANIDA Project ‘Support to Wild Life
Master Plan Implementation through the Improved Management of Totally Protected
Areas in Sarawak, Malaysia’ (SWMPI). The author is most grateful to Julian T. Inglis,
Chief Technical Advisor of the SWMPI Project, who has taken a lively interest in the
collection and description of these begonias; the paper would not have been written
without the inspiration and support provided by Dr Ruth Kiew, whose knowledge of
and interest in begonias prompted a closer look at the species occurring at Niah National
Park and who also checked the types at K and SING; and to Dr M.J.E. Coode for
translating the diagnoses into Latin, and to Dr A. Radcliffe-Smth for suggesting
‘stichochaete’ as the appropriate name for that species. The author also thanks the
staff of the Botany Unit, Forest Research Centre, and the Niah National Park, National
Parks and Wildlife Department, Sarawak Forest Department, for assistance during
collecting trips; the curator of SAR herbarium for permission to examine specimens;
and Joseph Pau for his excellent botanical illustrations.
Begonias from Niah National Park
Figure 5. Begonia subisensis
A Habit, B Juvenile shoot with variegated leaves, C Fruit. (A & C from $27574, B from
S78538).
87
88 Gard. Bull. Singapore 55 (2003)
Reference
Anderson, J.A.R. 1965. Limestone habitat in Sarawak. Symposium on Ecological
Research in the Humid Tropics. Kuching, Sarawak. Sarawak Government and
UNESCO. pp. 49-57.
Garden’s Bulletin Singapore 55 (2003) 89-95
A New Species of Curcuma L. (Zingiberaceae) from
Mizoram, India
J. SKORNICKOVA, M. SABU AND M.G. PRASANTHKUMAR
Department of Botany, University of Calicut,
673 635 Kerala, India.
Abstract
Curcuma rubrobracteata Skornickova, M.Sabu & Prasanthkumar sp. nov. is described from
Mizoram, India.
Introduction
Exploration for gingers in Mizoram, NE India, carried out by the authors for the
research project ‘Revision of Indian Zingiberaceae’ has brought an interesting species
of Curcuma to light.
The genus Curcuma L. naturally occurs in tropical and sub-tropical Asia with
a few species extending to Australia and the South Pacific, but economically important
or ornamental species are introduced elsewhere in the tropics. According to the latest
records, Curcuma has the largest number of species in India, Thailand, Indonesia and
China. The total number of species for this genus is still quite uncertain but is estimated
at around 100 species (Sirirugsa, 1996; Larsen et al., 1998). From India, 29 species
have so far been reported and accepted (Karthikeyan et al., 1989; Mangaly and Sabu,
1993). Several new species of Curcuma L. have been described in last 20 years from
India (Skornickova and Sabu, 2002). While working on Indian Zingiberaceae in August
and September 2002, the authors visited Mizoram, which is considered as the least
surveyed corner of India, where an undescribed species of the genus Curcuma L.
subg. Curcuma (Eucurcuma K. Schum.) was collected.
Curcuma rubrobracteata Skornickova, M. Sabu & Prasanthkumar, sp. nov.
Curcumae roscoeanae similis in coma inconspicua, spica c. 10 cm supra terram per
vaginam protrudenti, bracteis floriferibus rubris, floribus luteo-aurantiacis, calyce
trilobato, rhizomate repenti aromatico, lamina elliptico-lanceolata basi cuneata, infra
glabra, supra pubescenti in venis principalibus elevatis differt. Typus: India, Mizoram,
Lawngtlai District, on the way to Ngengpui Wildlife Sanctuary — Khomoi, 22° 30'N,
90 Gard. Bull. Singapore 55 (2003)
92° 46'E. 10.1X.2002. Skornickova & Prasanthkumar 86241 (holo MH; iso CALI, K,
SING).
Figure 1, Plate 1.
Rhizomatous herb, up to 1.5 m high. Rhizome creeping, slender, 10—30 cm long, c. 1 cm
diam., tan outside, scales triangular, papery, light brown, underground quickly
decaying, whitish yellow inside, aromatic, smell resembling Kaempferia galanga L.,
taste very bitter, sessile tubers absent, root tubers 2 x 1.5 cm, white inside, distanced
c. 5 cm away from main rhizome on 2 mm thick roots. Leafy shoot up to 1.5 m long,
leaves 4—6, pseudostem green, c. 40 cm long. Leaves petiolate, ligule 1 mm long,
light green, translucent, petiole green, glabrous 10-45 cm long (lower leaves with
shorter petioles). Lamina elliptic-lanceolate 35—60 x 10-16 cm, adaxially green,
pubescent along the raised veins, abaxially pale green, glabrous, base attenuate, tip
acuminate 2 cm long, midrib green, glabrous. /nflorescence terminal, but protruding
through the base of the pseudostem through lateral slits c. 3-10 cm above ground.
Peduncle 5—10 cm long, 7 mm diam., whitish, without vegetative bracts. Spike
10 x 7-9 cm. Coma absent. Bracts 20-26 per spike, all fertile, bright red or light red,
yellowish or yellowish green towards the base, bract 3.5 x 3.5 cm, glabrous, margin
0.3 mm hairy, hairs 0.1 mm long, subtending a cincinnus of 5—6 flowers, usually only
1—3 flowers per spike open at the same time. Bracteoles one per flower, 2—3.5 x 1-3 cm,
hyaline, glabrous, white, translucent with reddish dots on the apical part of the biggest
bracteoles. Flowers 6 cm long, yellow-orange, exserted, 1.5—2 cm longer than the
bracts. Calyx 1.2 cm, white, translucent, hyaline, glabrous, 3-toothed. Corolla tube
3.74 cm, light orange, glabrous. Corolla lobes light yellow-orange, dorsal lobe 14 x
8 mm fringed by red on upper mucronate portion; lateral lobes 12 x 7 mm. Labellum
15 x 17 mm, periphery yellow-orange, centre deep yellow-orange, obscurely 3-lobed,
middle lobe split about 2 mm. Lateral staminodes | x 1.1 cm; yellow-orange, hooded
over the anther. Stamen c. 9 mm long, anther versatile. Anther 6 x 2.5 mm, orange.
Anther spurs 3 mm long, incurved, orange. Anther thecae white, 5 mm long. Filament
yellow-orange, 5 mm long, constricted, 4 mm at base, 2 mm at upper part. Ovary
trilocular, 2 x 2.5 mm, pubescent with 0.6 mm long hairs; ovules many. Stigma white,
closely appressed within the anther lobes. Epigynous glands 2, pale orange, 4 mm
long, 0.9 mm diam. Fruits not seen.
Flowering: August — September.
Distribution: Hitherto known only from the type locality. From personal
communication with other specialists we suspect this species to be identical with
unidentified plants growing along the Thai-Burmese border.
91
Curcuma rubrobracteata
1
oe ee mee es ee
-—
i
Figure 1. Curcuma rubrobracteata
A. Habit; B. Inflorescence; C. Detail of leaf venation (adaxial side); D. Flower and subtending bract; E.
Bracteole; F. Part of flower showing lateral staminodes, dorsal corolla lobe and anther; G. Part of flower
showing labellum and anterior corolla lobes; H. Flower (side view); I. Calyx; J. Lateral staminode; K.
Anther (front); L. Anther (side); M. Stigma; N. Epigynous glands and ovary; O. Ovary (cross section).
Based on the type material Skornickova & Prasanthkumar 86241. del. J. Skornickova.
92 Gard. Bull. Singapore 55 (2003)
Habitat: Growing in undergrowth in teak plantations and along roadsides.
Etymology: This gorgeous species takes its name from the striking red bracts of its
inflorescence.
Other specimens examined: India, Mizoram, Lawngtlai District, on the way to
Ngengpui Sanctuary — Khomoi, 22° 30' N 92° 46' E, 10.[X.2002, Skornickova &
Prasanthkumar 86239 (CALI).
Notes: The most obvious characters of this species are the creeping rhizomes, bright
red bracts with no distinct coma and, in particular, the position of its inflorescence.
Inflorescences with no distinct coma are reported for a few species e.g. Curcuma
roscoeana Wall., C. albiflora Thwaites, C. ceratotheca K.Schum. and the recently
described C. rhomba J. Mood & K. Larsen, which also possesses a small rhizome
with linear growth (Mood and Larsen, 2001). The unique character of C.
rubrobracteata is the position of its inflorescence, which is so far unknown in the
genus Curcuma.
Many of the earlier taxonomists made attempts to classify the genus into
sections based on the position of the inflorescences. Roxburgh (1820) recognised
two sections based on the lateral or central inflorescence, while Horaninow (1862)
distinguished three sections, namely: I. Exanha (always lateral), I. Mesantha
(inflorescence invariably terminal), and III. Amphiantha (inflorescences both terminal
and lateral). However, the new species produces an inflorescence about 3—10 cm
above ground, which breaks through the pseudostem through lateral slits. This feature
has also been observed in the Zingiberaceae in the genus Plagiostachys Ridl., where
the inflorescence is borne on a peduncle and projects from the side of the leafy stalk
(Ridley, 1899). Smith (1985) pointed out that the inflorescence of Plagiostachys
species, although pushed out laterally, is actually terminal on the leaf shoot and so
shows affinity to the genus Alpinia Roxb., in which a few species, particularly A.
hansenit R.M.Smith and A. havilandii K.Schum., have similar apparently laterally
produced inflorescences.
Recently, Kress et al. (2002) carried out molecular studies on phylogeny and
classification of the family Zingiberaceae. The results based on analysis of DNA
sequences of the nuclear internal transcribed spacer (ITS) and plastid matK regions
suggest that the genus Curcuma, as it is accepted nowadays, is paraphyletic with
Plate 1. Curcuma rubrobracteata
1. Detail of flower and spike — seen from above; 2. Detail of flower and dorsal corolla
lobe — front view; 3. Detail of flower and bracts — lateral view; 4. Habit; 5. Detail of
spike protruding through the pseudostem; 6. Creeping rhizome and root tubers. All
photographs are of the type material Skornickova & Prasanthkumar 86241. Photo J.
Skornickova.
Curcuma rubrobracteata
mh
94 Gard. Bull. Singapore 55 (2003)
genera Hitchenia, Stahlianthus and Smithiatris, which also share cone-like
inflorescences of few flowered, congested bracts, and that genus Curcuma is itself
paraphyletic with three groups of species. Since the analysis was done with a limited
number of samples (six Curcuma species), the authors recommended that more species
of such large genus should be included in future molecular studies to shed more light
on the difficult question of generic boundaries and allied genera.
Acknowledgements
The authors thank the Department of Science and Technology, Govt. of India for
financial support (Order No. SP/SO/A-20/99 dt. 09.11.2001). We are also indebted to
staff of Forest Department of Mizoram for their hospitality and help to reach otherwise
inaccessible areas. The senior author thanks the Indian Council for Cultural Relations
New Delhi, India, and the Ministry of Education of the Czech Republic for awarding
a research fellowship, and to Singapore Botanic Gardens for providing facilities during
her repeated visits. We also thank Dr. J. F. Veldkamp, National Herbarium of
Netherlands, Leiden University Branch, and Dr. M. Svrcek, National Museum, Prague,
for the help with the Latin diagnosis.
References
Horaninow. 1862. Prodromus Monographiae Scitaminearum. Petropoli (St.
Petersburg), Russia.
Karthikeyan, S., S.K. Jain, M.P. Nayar and M. Sanjappa, 1989. Florae Indicae.
Enumeratio Monocotyledoneae. Botanical Survey of India, Calcutta.
Pp. 289-299.
Kress, J.W., L.M. Prince and K.J.Williams. 2002. The phylogeny and a new
classification of the gingers (Zingiberaceae): evidence from molecular data.
American Journal Botany. 89: 1682-1696.
Larsen, K., J.M.Lock, H. Maas and P.J.M. Maas. 1998. Zingiberaceae. In: K. Kubitzk1.
(Ed.) The Families and Genera of Vascular Plants. 4: 474-495.
Mangaly, J.K. and M. Sabu. 1993. A taxonomic revision of the South Indian species
of Curcuma L. (Zingiberaceae). Rheedea. 3: 139-171.
Mood, J. and K. Larsen. 2001. New Curcumas from South East Asia. The New
Plantsman. 8: 207—217.
Curcuma rubrobracteata 95
Ridley, H.N. 1899. The Scitamineae of the Malay Peninsula. Journal Straits Branch
Royal Asiatic Society. 32: 85-184.
Roxburgh, W. 1820. Monandria monogynia. Flora Indica. Serampore, India. Pp. 1-84.
Sirirugsa, P. 1996. The genus Curcuma of Thailand. In: T. L. Wu, Q. G. Wu and Z. Y.
Chen (Eds.) Proceedings 2" Symposium on Family Zingiberaceae. Zhongshan
University Press, China. Pp. 39-46.
Skornickova, J. and M. Sabu, 2002. The genus Curcuma L. in India: Resumé and
Future Prospects. In: A.P. Das (EIndiad.) Perspectives of Biology. Bishen Singh,
Mahendra Pal Singh, Dehradun, India Pp. 45-51.
Smith, R.M. 1985. A review of Bornean Zingiberaceae: 1 ( Alpinieae p.p.). Notes
Royal Botanic Garden Edinburgh. 42: 261-314.
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Garden’s Bulletin Singapore 55 (2003) 97-111
The Genetic Relations of Musa Species from Mount Jaya,
New Guinea, and a Reappraisal of the Sections of Musa
(Musaceae)
CAROL WONG!', GEORGE ARGENT?’, RUTH KIEW’, OHN SET?
AND YIK YUEN GAN!
'National Sciences Academic Group, National Institute of Education, Nanyang
Technological University, 1 Nanyang Walk, 637616 Singapore
"Royal Botanic Garden, Edinburgh, EH3 5LR, U.K.
3Singapore Botanic Gardens, | Cluny Road, 259569 Singapore
Abstract
Molecular analysis using amplified fragment length polymorphism (AFLP) confirms the status
of Musa banksii F. Muell. as a subspecies of M. acuminata Colla and shows M. johnsii Argent
to be genetically more similar to the Australian M. jackeyi W. Hill than to M. lolodensis
Cheesman from New Guinea. In addition, AFLP analysis supports only two sections as
genetically distinct, namely Sect. Musa (including Sect. Rhodochlamys) and Sect. Callimusa
(including Sect. Australimusa). No material for Sect. [gentimusa was available for study.
However, for practical purposes of grouping banana species, four informal groups are recognised
— the ‘acuminata’ and ‘ornata’ groups within Sect. Musa, and the ‘coccinea’ and ‘textilis’
groups within sect. Callimusa. A key to the sections and informal groups is provided.
Introduction
Wild bananas are found throughout Asia and Malesia extending into Australia and
the Pacific. They have become an increasingly conspicuous element of the vegetation
as they invade forest margins along logging roads and openings in forest. As roads
penetrate deeper into forests, new banana species are coming to light.
Much still remains to be discovered about the relationships between wild
banana species, which have been grouped into five sections (Cheesman, 1947; Argent,
1976) based on chromosome number and morphology. However, some of the species
recently described do not fit comfortably into these sections (Argent, 2000, 2001),
which calls into question their taxonomic validity. Also, molecular studies indicate
that not all the sections are genetically uniform and distinct from each other (Jarret &
98 Gard. Bull. Singapore 55 (2003)
Gawel, 1995; Wong et al., 2002).
Bananas present a challenge to the collector who wishes to turn them into
herbarium specimens, which unless accompanied by spirit material, detailed field
notes and colour pictures, are often almost worthless. Molecular techniques are
therefore particularly useful in this genus and have been used to study genetic diversity
at the subspecific (Wong et al., 2001a), specific (Wong et al., 2001b) and sectional
levels (Jarret & Gawel, 1995, Wong et al., 2002). AFLP is a DNA fingerprinting
technique, which was developed by Vos et al., (1995). It is based on selective PCR
amplification of DNA restriction fragments under stringent conditions. It can be used
for DNA of any origin and complexity and is reported to be both reproducible and
reliable (Vos et al., 1995). AFLP combines the reliability of RFLP with the power of
PRC.
In this study, AFLP (amplified fragment length polymorphism) was used to
analyse the genetic relations of two species collected from Mt Jaya, Papua (formerly
Irian Jaya), New Guinea: (a) to study the genetic relationship of Musa johnsii Argent,
which is unique in the genus in having a fruit with a sterile mucilaginous pith chamber
that occupies the distal third of the fruit (Argent, 2001); and (b) to reassess the status
of M. banksii F. Muell., which was reduced to a subspecies of M. acuminata Colla by
Simmonds (1956) but is still considered to be a distinct species by Argent (1976).
In addition, the taxonomic status of the five sections of Musa is reviewed in
the light of molecular studies.
Materials and Methods
Plant Materials
Twelve samples were used for the present study (Table 1). These included Musa
species from sections Musa, Rhodochlamys, Callimusa and Australimusa and a species
of the related genus Ensete as an outgroup taxon. Voucher specimens were deposited
in the herbaria of the Singapore Botanic Gardens and the Royal Botanic Garden
Edinburgh.
DNA Extraction
Leaf tissue was used for AFLP analysis and prepared using a procedure from Zhang
et al. (1997). Plant DNA was extracted using the CTAB method according to Reichardt
and Rogers (1993) as outlined in Wong et al. (2001b).
AFLP analysis
The AFLP analysis was carried out according to Vos et al. (1995) with minor
modifications. Restriction digests of genomic DNA with EcoRI and MselI were carried
out at 37 °C for | h. Following heat inactivation of the restriction endonucleases, the
genomic DNA fragments were ligated to EcoRI and MselI adapters overnight at 16°C
99
Musa in New Guinea & Section of Musa
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Gard. Bull. Singapore 55 (2003)
to generate template DNA for amplification. PCR was performed in two consecutive
reactions. The template DNA generated was first pre-amplified using AFLP primers
each having one selective nucleotide. The PCR products of the pre-amplification
reaction were then used as template after dilution 5-fold in sterile water, for selective
amplification using two AFLP primers, each containing three selective nucleotides.
A total of eight primer combinations were used in this study (Table 2). The final PCR
products were run on a 6% denaturing polyacrylamide gel in 1X TBE buffer. The
EcoRI primers used were not radioactively labelled as in the original protocol. Instead,
a modified silver staining method was used (Loh et al., 1999).
Table 2 The sequence of primers used in the AFLP analysis.
Name / Abbreviation
GYY 101/ EA+
GYY 102/ EA-
GYY 103/ MA+
GYY 104/ MA-
GYY 105/E-A
GYY 107/ E-AAC
GYY 108/ E-AAG
GYY 109/ E-ACA
GYY 110/ E-ACT
GYY 111/ E-ACC
GYY 112/ E-ACG
GYY 113/ E-AGC
GYY 114/ E-AGG
GYY 106/ M-C
GYY 115/ M-CAA
GYY 116/ M-CAC
GYY 117/ M-CAG
GYY 118/ M-CAT
GYY 119/ M-CTA
GYY 120/ M-CTC
GYY 121/M-CTG
GYY 122/ M-CTT
Enzyme
EcoRI
EcoRI
Msel
Msel
EcoRI
EcoRI
EcoRI
EcoRI
EcoRI
EcoRI
EcoRI
EcoRI
EcoRI
Msel
Msel
Msel
Msel
Msel
Msel
Msel
Msel
Msel
Type
Adapter +
Adapter -
Adapter +
Adapter -
Primer +1
Primer +3
Primer +3
Primer +3
Primer +3
Primer +3
Primer +3
Primer +3
Primer +3
Primer +1
Primer +3
Primer +3
Primer +3
Primer +3
Primer +3
Primer +3
Primer +3
Primer +3
Sequence (5’-3’)
CTCGTAGACTGCGTACC
AATTGGTACGCAGTCTAC
GACGATGAGTCCTGAG
TACTCAGGACTCAT
GACTGCGTACCAATTCA
GACTGCGTACCAATTCAAC
GACTGCGTACCAATTCAAG
GACTGCGTACCAATTCACA
GACTGCGTACCAATTCACT
GACTGCGTACCAATTCACC
GACTGCGTACCAATTCACG
GACTGCGTACCAATTCAGC
GACTGCGTACCAATTCAGG
GATGAGTCCTGAGTAAC
GATGAGTCCTGAGTAACAA
GATGAGTCCTGAGTAACAC
GATGAGTCCTGAGTAACAG
GATGAGTCCTGAGTAACAT
GATGAGTCCTGAGTAACTA
GATGAGTCCTGAGTAACTC
GATGAGTCCTGAGTAACTG
GATGAGTCCTGAGTAACTT
Musa in New Guinea & Section of Musa 101
Data analysis
For the diversity analysis, bands were scored as present (1) or absent (0) to form a raw
data matrix. A square symmetric matrix of similarity was then obtained using Jaccard’s
Similarity Coefficient Coefficient [x/(y-z)], where x is the number of fragments in
common between two taxa, y is the total number of fragments scored, z is the number
of fragments absent in both taxa, from the raw data matrix. Genetic diversity estimates
(GDEs) were then calculated as 1 minus Jaccard’s Similarity Coefficient and used for
cluster analysis using the Unweighted Pair Group Method with Arithmetic mean
(UPGMA) technique of the NEIGHBOR program in PHYLIP version 3.5c
(Felsenstein, 1993). The dendrogram was drawn using TREEVIEW version 1.6.1
(Page, 1996).
Results
AFLP Profiles
The AFLP profiles generated using eight primer combinations contained highly
informative bands, which distinguished all taxa examined and provided valuable
information on genetic relationships. Figure | illustrates an AFLP profile obtained.
Only unambiguous bands of size 50-500 base pairs were scored. A total of 201 bands
were analysed out of which (95%) were polymorphic across all samples examined.
On average, 27 bands were scored per primer pair.
Fifty-nine unique bands were observed for the taxa examined (Table 3). All
species except for Musa textilis were characterised by unique markers ranging from 1
to 18. M. johnsii is characterised by six unique bands, confirming it is a distinct species.
M. banksii is characterised by four unique bands. The genetic markers observed will
be useful for the development of probes in Musa breeding programmes. The genetic
diversity estimates are shown in Table 4.
Discussion
Position of Musa johnsii
This species has strikingly distinct morphological characters, such as the compact,
subglobose head of schizocarpic fruits that are unique in the distal third being filled
with pale pinkish orange, mucilaginous pith (Argent, 2001). AFLP analysis confirms
it is a distinct species (Fig. 2). Argent (2001) suggested it was closely related to the
schizocarpic M. lolodensis as the seeds of the two species are remarkably similar but
Table 4 and Fig. 2 show it is genetically more similar to the Australian M. jackeyi than
102 Gard. Bull. Singapore 55 (2003)
it is to the New Guinea M. lolodensis. All three species belong in Sect. Callimusa in
the ‘textilis’ group (see below). This is another example where conspicuous
morphological differences do not reflect genetic similarity (Jarret & Gawel, 1995).
Status of Musa banksii
Simmonds (1956) reduced this species to a subspecies of Musa acuminata based on
experiment and field observations. However, Argent (1976) maintained it as a distinct
species as it did not hybridise with M. acuminata subsp. malaccensis when they were
grown together for many years in the Lae Botanic Garden, Papua New Guinea. This
taxon also differs from the other M. acuminata subspecies in its female flowers having
some fertile stemens, in the non-imbricating bracts in the male bud, and in producing
a very large number of seeds (up to 400) as compared with 40-50 in subspecies from
the lowlands of Peninsular Malaysia and Thailand (Simmonds, 1995).
Results of several studies using molecular techniques are now available. Gawel
& Jarret (1991) used cpDNA RFLP, which generated a phenogram with M. banksii
embedded among the other subspecies of M. acuminata studied, indicating that it is
not genetically distinct at the species level. Jarret & Gawel (1995) using total-DNA
RFLP again found that M. banksii clustered with the other M. acuminata subspecies
in their phenogram. Our study using AFLP showed M. banksii clustering with the
other subspecies of M. acuminata (Fig. 2), again indicating that genetically it is not a
distinct species and that subspecific rank would be more appropriate. However, the
genetic diversity estimates (Table 4) indicate it is the least similar among the four
subspecies studied, perhaps a reflection of a combination of its unique characters
listed above and its most south-easterly distribution of all M. acuminata subspecies.
Sections and groupings of wild banana species
Cheesman (1947) created four sections within the genus Musa - sections Eumusa
(now Musa), Rhodochlamys, Callimusa and Australimusa - as a convenient way of
grouping the species. He noted that the sections were not of equal rank and that in
some characters sect. Australimusa was intermediate between sect. Callimusa and
sect. Musa, and that the division between sect. Musa and sect. Rhodochlamys was
‘unessential’. He speculated that sect. Musa and sect. Australimusa were the earliest
to diverge and that sect. Rhodochlamys was an offshoot from sect. Musa. He was in
two minds as to whether sect. Callimusa had diverged directly from sect. Australimusa
or, as he considered the barrel-shaped seeds with a large oil/air space a very significant
character, might be an earlier divergence from sect. Musa. Argent (1976a) described
a fifth section, /gentimusa, with the single species M. ingens N.W. Simmonds.
Subsequent to Cheesman’s work, new species have been described that do
not conform to his concepts of the sections so that the distinction between the sections
is becoming blurred. Molecular techniques have opened a new avenue of enquiry.
Studies carried out so far (Jarret & Gawel, 1995; Wong et al., 2002, and this study)
Musa in New Guinea & Section of Musa 103
Miz3 45 67 8°910l1 12
at =e
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Figure 1. AFLP profile generated using primer pair 1 (E-AAC, M-CAA). Lane M: pUC19/Hpall
molecular weight marker, lane 1: Musa johnsii, lane 2: Ensete superbum, lane 3: M. lolodensis, lane 4:
M. textilis, lane 5: M. jackeyi, lane 6: M. borneensis, lane 7: M. violascens, lane 8: M. banksii, lane 9: M.
acuminata ssp. siamea, lane 10: M. acuminata ssp. truncata, lane 11: M. acuminata ssp. microcarpa,
lane 12: M. sanguinea.
Gard. Bull. Singapore 55 (2003)
104
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105
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106 Gard. Bull. Singapore 55 (2003)
Ensete superbum
Musa lolodensis
Musa violascens
Musa textilis
Musa johnsii
Musa jackeyi
Musa borneensis
Musa acuminata ssp. banksii
Musa acuminata ssp. truncata
Musa acuminata ssp. siamea
Musa acuminata ssp. microcarpa
Musa sanguinea
0.1
Figure 2. UPGMA cluster analysis of AFLP data generated by eight primer combinations for 12 taxa of
Musa depicting patterns of genetic diversity. Scale depicts genetic diversity estimates (GDEs).
Musa in New Guinea & Section of Musa 107
strongly support two main groups within the genus Musa, which correlate with
chromosome number, i.e. those with a somatic number of x=11 (sect. Musa and sect.
Rhodochlamys) and x=10 (sect. Callimusa and sect. Australimusa, including M.
beccarii with a somatic number of n=9). On the other hand, they also show that sect.
Rhodochlamys is not genetically distinct from sect. Musa, and that sect. Australimusa
is not distinct from sect. Callimusa. No material of Musa ingens (sect. [gentimusa,
somatic no. x=7) has been available for molecular analysis.
The conspicuous distinction between sect. Musa and sect. Rhodochlamys is
the tall pseudostem and pendulous inflorescences with dull coloured bracts, which
mostly become revolute on fading, of the former, compared with the short pseudostems
and erect inflorescences and brightly coloured bracts that do not become revolute of
the latter. Apart from pseudostem height, these characters are related to the pollination
syndrome: the flowers on pendulous inflorescences being pollinated by nectiferous
bats and the erect ones by nectiferous birds (sunbirds and spiderhunters). Molecular
studies show that many species classed within sect. Rhodochlamys are in fact more
closely related to species within sect. Musa than they are to other species in sect.
Rhodochlamys, which indicates that bird-pollination in bananas has evolved
independently several times. Simmonds (1962) considered bat pollination ancestral
to bird pollination in the genus Musa. Both Simmonds (1962) and Shepherd (1999)
suggested that these two sections be amalgamated. Molecular studies (Jarret & Gawel,
1995; Wong et al., 2002 and this study) strongly support this view.
Cheesman (1947) was most impressed with the distinct barrel-shaped seeds
with the large oil/air space of sect. Callimusa, so much so that he thought it would
have warranted generic status had there not been species in sect. Australimusa that
shared characters of both sect. Musa and sect. Callimusa. However, as more species
have been discovered, seed morphology has been shown to be much more variable
(Argent, 2000). In general, sect. Ca/limusa includes species with erect inflorescences
with colourful bracts compared with most species in sect. Australimusa with pendent
inflorescences (although there are exceptions in both sections), again indicating that
the erect inflorescence with bird-pollinated flowers has evolved several times.
Molecular studies (Wong et a/., 2002; and this study) also illustrate that there is no
genetic justification for keeping them separate as species in sect. Callimusa cluster
among species of sect. Australimusa (Fig. 2).
To put banana taxonomy on a firmer basis where the sections reflect genetic
similarity among the species, three sections are recognised here:
Section Musa (Baker) Cheesman, Kew Bull. (1948) 108.
Basinym: subgen. Eumusa Baker, Ann. Bot, 7 (1893) 205, 208.
Type: M. paradisiaca L.
Syn: Sect. Rhodochlamys Baker, Ann. Bot. 7 (1893) 205.
Type: M. ornata Roxb.
108 Gard. Bull. Singapore 55 (2003)
Section Callimusa Cheesman, Kew Bull (1947) 112.
Type: M. coccinea Andrews
Syn. Sect. Australimusa Cheesman, Kew Bull. (1947) 110.
Type: M. textilis Nee
Section Ingentimusa Argent, Notes Roy. Bot. Gard. Edinb. 35 (1976) 111.
Type: M. ingens N.W. Simmonds.
However, Cheesman’s point that his four sections are a convenient way of
grouping bananas still holds as the bird-pollinated species with colourful bracts in
sect. Rhodochlamys and sect. Callimusa are important horticulturally, while the edible
bananas fall into the other two groups, sect. Musa with the M. acuminata x M. balbisana
hybrids and sect. Aastralimusa, which includes the Fe’i cultivar group. In addition, as
Cheesman remarked, some groups are easily identified in the field. For example,
most species in the ‘coccinea’ group are readily recognised by the pronounced scarious
auricle along the leaf shoulder (Argent, 1976). For these reasons,we suggest that an
informal grouping be adopted to satisfy this need and, to avoid confusion with the
sections, should be named for the ‘type’ species for each group. Thus, sect. Musa
would include the ‘acuminata’ and ‘ornata’ groups and sect. Callimusa the “coccinea’
and ‘textilis’ groups.
Jarret & Gawel (1995) considered M. coccinea Andrews sufficiently distinct
in their molecular study to be placed in a group of its own, though it should be noted
that this result could be due to the fact that no other species in the ‘coccinea’ group
was included in their study. In fact it differs from the rest of the group morphologically
in lacking the pronounced rugose auricles but when other species in the ‘coccinea’
group are included, M. coccinea clusters with them (Wong ef al., 2002). In any case,
if this argument is followed, it would end up with each of the unusual or anomalous
species, such as M. johnsii or M. suratii Argent, each being placed in a group of their
own, which defeats the purpose of grouping species in the first place.
To show the diagnostic characters of each of these groups, a key to the sections
and informal groups is given below:
Key to Banana Groups
(based on Cheeseman, 1947, and Argent, 1976)
la. Section Musa. Chromosome number x=11. Bracts usually more or less sulcate,
often more or less glaucous, rarely or never polished, convolute or more or less
imbricate in the bud, usually strongly revolute on fading. Seeds often dorsiventrally
COMMTCSSER, SOIICLMIES SIO OOS Eo sce eosuiccie ease paces DCA as kee 2
Musa in New Guinea & Section of Musa 109
2a. Pseudostems commonly exceeding 3 m tall. Inflorescences pendent or
semi-pendent, fruits reflexing towards the base of the rachis. Bracts commonly
dull-coloured, green, brownish or dull purple. Flowers many to a bract, in
OO Ss, Rk Pe TE A se acuminata group
2b. Pseudostems commonly less than 3 m tall. Inflorescences erect, fruits
pointing towards the apex of rachis. Bracts brightly coloured, often pink or
red. Flowers few to a bract, usually in a single series............... ornata group
1b. Bracts plane, firm in texture, rarely or never glaucous, strongly imbricate in
bud, polished or not, usually not revolute on fading. Seeds barrel-shaped or
SE smee ty EG LZ, DANS ass.) GD, Sa, a A... J
3a. Section Ingentimusa. Chromosome number x=7. Pseudostems
c.15 m tall. Leaf shoulders reflexed, non-scarious. Inflorescences
pendent. Bracts not polished. Seeds strongly angled, 8—9 mm diam.
3b. Section Callimusa Chromosome number x=10. Pseudostems
short or tall. Leaf shoulders scarious or non-scarious, never reflexed.
Inflorescences erect or pendent. Bracts polished on the outside. Seeds
barrel-shaped or strongly angled, up to 5 mm diam.................... 4
4a. Pseudostems commonly less than 3 m tall and
inflorescences erect or sometimes pseudostems more than 3 m
tall and inflorescences pendent (e.g. M. borneensis). Leaf
shoulders usually with well-developed scarious auricle,
rarely absent (e.g. M. coccinea). Seed often barrel-shaped or
cylindrical with a well-developed oil/air space, rarely
Sek a ice nee ada sibs sdaw rotates coccinea group
4b. Pseudostems more than 3 m tall. Inflorescence commonly
pendent, sometimes erect (e.g. M. maclayi). Leaf shoulders
commonly without a scarious auricle, sometimes scarious
(e.g. M. lolodensis). Seeds subglobose or dorsiventrally
compressed, sharply angled, without a well-developed oi1/
Bina pane cotati ah OWL AASTL.WOee!.. textilis group
Acknowledgements
We are indebted to the Directors of the Royal Botanic Garden Edinburgh and the
Singapore Botanic Gardens for permission to collect leaf samples from their living
collections; to the Academic Research Fund, National Institute of Education, Nanyang
Technological University, for research grant RP 12/98/GYY; and GA thanks R.J.
Johns, T. Utteridge and the Royal Botanic Gardens Kew for making possible field
work on Mt Jaya.
110 Gard. Bull. Singapore 55 (2003)
References
Argent, G. 1976. The wild bananas of Papua New Guinea. Notes Royal Botanic Garden
Edinburgh. 35: 77-114.
Argent, G. 2000. Two interesting wild Musa species (Musaceae) from Sabah, Malaysia.
Gardens’ Bulletin Singapore. 52: 203-210.
Argent, G. 2001. Contributions to the flora of Mount Jaya VI. A new banana, Musa
Johnsii (Musaceae) from New Guinea. Gardens’ Bulletin Singapore. 53: 1-7.
Cheesman, E.E. 1947. Classification of the bananas. III. The genus Musa L. Kew
Bulletin. 106-117.
Felsenstein, J. 1993. PHYLIP (Phylogeny Inference Package) version 3.5c. Distributed
by the author. Department of Genetics, University of Washington, Seattle.
Gawel, N.J. and R.L. Jarret. 1991. Choroplast DNA restriction fragment length
polymorphism (RFLPs) in Musa species. Theoretical and Applied Genetics. 81:
783-786.
Jarret, R.L. and N.J. Gawel. 1995. Molecular markers, genetic diversity and systematics
in Musa. In: Gowen, S. (ed.). Bananas and Plantains. Chapman & Hall, U.K.
Chapter 3.
Loh, J.P, R. Kiew, A. Kee, L.H. Gan and Y.Y. Gan. 1999. Amplified fragment length
polymorphism (AFLP) provides molecular markers for the identification of
Caladium bicolor cultivars. Annals of Botany. 84: 155-161.
Page, R.D.M. 1996. TREEVIEW: An application to display phylogenetic trees on
personal computers. Computer Applications in the Biosciences. 12: 357-358.
Reichardt, M.J. and S.J. Rogers. 1993. Plant DNA isolation using CTAB. In: FM.
Ausubel, R. Brent, R.E. Kingston, D.D. Moore, J.G. Seidman, J.A. Smith, and K.
Struhl (eds). Current Protocols in Molecular Biology. Supplement 22, John Wiley
and Sons, USA.
Shepherd, K. 1999. Cytogenetics of the Genus Musa. International Network for the
Improvement of Banana and Plaintain, Montpellier, France.
Simmonds, N.W. 1956. Botanical results of the Banana Collecting Expedition, 1954-5.
Kew Bulletin. 463-489.
Musa in New Guinea & Section of Musa Lil
Simmonds, N.W. 1962. The Evolution of Bananas. Longmans, U.K.
Vos, P., R. Hogers, M. Bleeker, M. Reijans, T. van de Lee, M. Hornes, A. Frijters, J.
Pot, J. Peleman, M. Kupier and M. Zabeau. 1995. AFLP: a new technique for
DNA fingerprinting. Nucleic Acids Research. 23: 4407-4414.
Wong, C., R. Kiew, J.P. Loh, L.H. Gan, S.K. Lee, Ohn Set, S. Lum, Y.Y. Gan. 2001a.
Genetic diversity of the wild bananas Musa acuminata Colla in Malaysia as
evidenced by AFLP. Annals of Botany. 88: 1017-1025.
Wong, C., R. Kiew, Ohn Set, A. Lamb, S.K. Lee, L.H. Gan, Y.Y. Gan. 2001b. Sectional
placement of three Bornean species of Musa (Musaceae) based on AFLP. Gardens’
Bulletin Singapore. 53: 327-341.
Wong, C., R. Kiew, G. Argent, Ohn Set, S.K. Lee, Y.Y. Gan. 2002. Assessment of the
validity of the sections of Musa (Musaceae) using AFLP. Annals of Botany. 90:
231-238.
Zhang, W.P, J.F. Wendel and L.G. Clark. 1997. Bamboozled again! Inadvertent
isolation of fungal rDNA sequences from bamboos (Poaceae: Bambusoideae).
Molecular Phylogenetics and Evolution. 8: 205-217.
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Garden’s Bulletin Singapore 55 (2003) 113-123
Begonias from the Bau Limestone, Borneo,
including a New Species
RUTH KIEW' AND CONNIE GERI?
'The Herbarium, Singapore Botanic Gardens, Singapore
* The Sarawak Biodiversity Centre, Kuching, Sarawak, Malaysia
Abstract
Six Begonia species (Begoniaceae) are found on limestone hills in the Bau area, Kuching,
Sarawak. One, a new species, Begonia lailana Kiew & Geri, is described. A key and habitat
notes for all species are provided. The male and female flowers of B. calcarea Ridl. and the
female flowers of B. congesta Ridl., not previously recorded, are here described for the first
time.
Introduction
In 1906, Ridley published a paper on the begonias of Borneo that included twenty
species, of which five were new species from the Bau limestone (1° 25'N 110° 9'E),
35 km from Kuching, Sarawak. Most were based on his own collections. Two of
these were imperfectly known so that descriptions of the male and female flowers of
Begonia calcarea Ridl. and the female flowers of B. congesta Ridl. were lacking.
As part of the recent biodiversity survey of the Bau limestone carried out by
the Sarawak Biodiversity Centre, we collected specimens of begonias from 20 hills.
All the species described by Ridley were recollected with the addition of a new species,
which is described in this paper. Flowering material of B. calcarea was also collected
and the flowers are described below.
In addition, the habitats of the species, which are quite different, were recorded.
The diversity of begonias on a single limestone hill can be attributed to individual
species occupying different and non-overlapping niches (Kiew, 1998). Most of the
hills at Bau are home to at least three species, B. pendula Ridl., B. rubida Ridl. and B.
speluncae Ridl., with four species collected from Gunung (G.) Doya and G. Jebong
and five from G. Kawa. Begonia rubida grows near the summit of limestone hills
above the tree canopy exposed to sunlight or in light shade, while B. pendula grows
below the tree canopy on damp, mossy, vertical shaded rock-faces. B. speluncae, as
114 Gard. Bull. Singapore 55 (2003)
its name suggests, is common around cave mouths but also grows on dry rock-faces
near the base of the hills where it occurs. The less common species include B. congesta,
which grows on limestone boulders or damp low rock-faces near the cliff base, and B.
calcarea and the new species, B. lailana Kiew & Geri, which both grow close to
limestone hills on soil associated with the limestone. This soil is frequently more
friable and more free-draining than soil derived from other rock types and often
supports a different suite of plant species. It is significant that these last three species
are less common than the others as this habitat is more susceptible to disturbance,
particularly from agriculture, which encroaches towards the base of the hills.
Several of these species are decorative and have ornamental value. Begonia
calcarea is a handsome species with broad upstanding leaves, which are covered in
striking long magenta hairs, particularly on the petiole. In addition, it has large, bright
orange or orangey-red flowers with a wavy margin. B. pendula has attractive foliage
—its leaves are variegated with silver-green spots. Ridley (1906) described B. speluncae
as ‘one of the most attractive limestone species’ with its tufts of small round leaves.
For identifying species, Ridley’s key is not entirely satisfactory as both B.
pendula and B. rubida key out under ‘stems short, erect and leafy’. B. pendula is a
creeping and often pendulous species and B. rubida has tall stems. A key to the Bau
limestone species is therefore provided.
Key to the Bau Limestone Begonias
la. Stems erect and cane-like 2
1b. Stems rhizomatous or creeping ~
2a. Leaves with veins prominently forked and narrowly diverging,
fruits oblong 2. B. congesta
2a. Leaves with veins widely diverging when they fork, fruit much wider
distally 3
3a. Leaves ovate, petiole at an angle to the midrib 5. B. rubida
3b. Leaves oblanceolate, petiole straight with the midrib 3. B. lailana
4a. Stems thin and creeping, leaves widely spaced, longer than broad,
spotted silver-green 4. B. pendula
4b. Stem thick and rhizomatous, leaves tufted, as wide as broad, dark
green, not spotted 5
5a. Robust plant, lamina more than 15 cm wide, petiole more than 15 cm
long with dense long magenta hairs, flowers orange 1. B. calcarea
Begonias from Bau, Borneo 15
5b. Diminutive plant, lamina up to 4 cm wide, petiole up to 7 cm long, hairs not
long nor magenta, flowers white 6. B. speluncae
1. Begonia calcarea Ridl., J. Str. Br. Roy. As. Soc. 46 (1906) 260. Holotype:
Sarawak, Gunung Bra’ang Haviland s.n. (not seen).
Sect. Diploclinium
Ridley (1906) described this handsome begonia from a fruiting specimen. It is a very
striking plant due not only to its broad, upstanding leaves but also to its long magenta
hairs, which cover the whole plant and are particularly striking on the petioles. It is
unusual among Bornean begonias in having orange flowers (most begonias have pale
pink or white ones). The flowers are produced below the leaves from the prostrate
rhizome. They are described here from SBC 3854 for the first time.
Inflorescences from the leaf axils or from the prostrate rhizome. Female flowers, 1—2
at one node, bracts c. 10 x 4 mm. Male flowers in umbels 2.5—5 cm long, rachis rosy
pink to pale pink, male flowers 3-6; bracts lanceolate 7-10 x 2-4 mm, apex setose;
bracteoles 2—6 x 1-3 mm. Male flowers with pedicel 4-19 mm long, tepals 4, orangey-
red to deeper red around the margin, margin undulate, glabrous inside and out;
isomorphic, outer two broadly oval to almost rotund, apex rounded, 7—11 x 5-9 mm,
inner ones slightly narrower, 7—11 x 4-6 mm; stamen cluster hemispherical, c. 4-5 x 8 mm,
almost sessile, stamens 25-35, filament c. 1.5—2 mm long, anthers deep yellow,
obovate, c. 1.5—2 mm long, apex emarginate. Female flowers with rosy pink peduncle
and pedicel, bracteole oval, c. 5 x 2 mm, apex acuminate, ovary c. 19 x 20 mm,
glabrous, locules 3, placentas bifid; tepals 5, isomorphic, orangey-red, margin undulate,
c.6x 7mm, stigma and styles c. 4 mm long. Fruits with peduncle and pedicel 20-40 mm
long, capsule 11—21 x 15-21 mm, wings 3, equal, rounded, 4-12 x 5—7 mm, thinly
fibrous. Seeds barrel-shaped, light brown, c. 0.2 mm long, collar cells (0.25—)0.75 length
of seed.
Doorenbos et al. (1998) doubtfully included Begonia calcarea in sect.
Diploclinium. Examination of the female flower shows its ovary and fruit to have
three locules and bifid placentas, which confirms its position within this section.
Haviland recorded its habitat as ‘top of Mt Bra’ang, old jungle, limestone’,
hence the name ‘calcarea’ that Ridley gave. However until our survey, no further
specimens had been identified as this species so that there was some doubt that it was
a limestone plant. It is certainly a rare species as, although 20 hills were surveyed, it
was only found at the base of two of them. It is the most endangered among the Bau
limestone species because its habitat is vulnerable to encroaching agricultural practices
116 Gard. Bull. Singapore 55 (2003)
from farms close to the base of the hills.
A similar specimen (Native Collector 1255) was collected from Gunung Dulit
casting doubt as to whether this species is restricted to limestone and endemic to
limestone in the Kuching District.
Distribution: Malaysia, Sarawak - Kuching limestone and possibly also from non-
limestone habitats on G. Dulit.
Specimens examined: Gunung (G.) Angob Anderson §27513 (SAR); G. Kawa Connie
Geri et al. SBC 3577 (SBC); G. Lanyang (G. Meraja) Burtt B8/54 (SAR), G. Manok
Burtt B8134 (SAR), B8154 (SAR); G. Tabai Connie Geri et al. SBC 3854 (SBC).
2. Begonia congesta Ridl., J. Str. Br. Roy. As. Soc. 46 (1906) 253. Holotype:
Sarawak, Bau Ridley s.n. (not seen).
Sect. Petermannia
This is a distinct begonia because its leaves are corrugate due to the many, narrowly
bifurcating veins. As its name suggests, the inflorescence appears congested because
the lateral branches are clustered at the base but above which there is a long (up to 14
cm long) rachis bearing male flowers. This rachis dies and falls before the fruits are
mature and was obviously not present in the plants Ridley described. The plant is
protogynous with two female flowers produced at the base followed by many small
male flowers, which are produced later on short lateral branches and on the erect
rachis.
Male flowers with pedicel 5—7 mm long, with pale brown hairs c. 0.25 mm long;
tepals 2, cream and rosy pink at base, minutely hairy outside, glabrous inside, upper
rotund, 4-5 x 4—5 mm, lower broadly oval, 4-5 x 3—4 mm; stamen cluster
hemispherical, c. 3—3.5 x 3-4 mm, joined at base in a stalk c. 0.5 mm long; stamens c.
35, filament c. 0.75 mm long, anther obovate, c. | mm long, apex not emarginate.
Female flowers with pedicel 4-11 mm long with dense glandular hairs, ovary oblong,
to 20 x 17 mm, wing 3-4 mm wide, isomorphic, rounded distally, locules and wings
minutely hairy, locules 3, placentas bifid; tepals 5, isomorphic, oval, c. 18 x 6 mm,
toothed in distal half, each tooth terminating in a hair, minutely hairy outside, glabrous
inside; styles 3, c. 3 mm long, bifid, stigma a spiral papillose band. Fruits pendent,
broadly oblong, to 2.5 x 1.7 cm, wings 3, equal, thin and fibrous. Seeds barrel-shaped,
c. 0.3 mm long, collar cells c. 0.75 seed length.
Ridley (1906) recorded the tepals as shorter than the stamens (surely an error
as the stamens are less than 2 mm long) and the stamen number as 12. We counted
Begonias from Bau, Borneo cig
about 35 stamens.
The species is endemic in the Kuching limestone. Ridley cited a specimen
from the Niah limestone under this species but this has proved to be a distinct species
(Pearce, 2003).
Distribution: Malaysia, Sarawak - endemic in the Kuching limestone.
Specimens examined: Bau Abang Mohtar S52906 (SAR), Brooke 9875 (SING),
Purseglove P4468 (SAR, SING); Bidi Mamit S42147 (SAR); Bukit Kapur Burtt &
Woods B188&1 (SAR); Bukit Krian Anderson $25137 (SAR); Fairy Cave Jugah et al.
$70076 (SAR); G. Angob Anderson $27493 (SING); G. Batu Martin S39276 (SAR),
Meekiong et al. SBC 2125 (SBC); G. Doya Julia et al. SBC 2062 (SBC), Meekiong et
al. SBC 2244 (SBC), Raymond et al. SBC 3459 (SBC); G. Jebong Meekiong et al.
SBC 2076 (SBC), Paul & Ilias $25626 (SAR); G. Kawa George S38280 (SAR),
Meekiong et al. SBC 3113 (SBC); G. Krian Meekiong et al. SBC 1614 (SBC); G.
Lanyang Julia et al. SBC 1224 (SBC), SBC 2938 (SBC); G. Pambur Julia et al. SBC
1270 (SBC); G. Ropith Donny et al. SBC 2001 (SBC), Julia et al. SBC 1295 (SBC);
Seburan Anderson 8984 (SAR), S/4580 (SAR), Woods B2890 (SAR); G. Tai Ton
Denis et al. SBC 1410 (SBC), Julia et al. SBC 726 (SBC); G. Tongga Burtt B&179
(SAR).
3. Begonia lailana Kiew & Geri, sp. nov.
Differt a Begonia oblongifolia folis latioribus (8—10 cm nec 5 cm latis), floribus nec
albis et fructis alis latioribus (8—11 mm nec 5 mm latis).
Typus: Malaysia, Sarawak, Bau, Gunung Kawa. Connie Geri et al. SBC 3753 (holo
SAR, iso K, L, SAN, SBC, SING).
Section: Petermannia
Figure 1
Cane-like begonia up to | m tall with several erect stems from the base, some plants
branching near the base to form a bushy crown. Stem succulent becoming woody,
reddish or greenish brown, swollen at nodes, up to c. 7 mm diam., glossy and glabrous
or hispid with uniseriate translucent hairs c. 2 mm long, dense on the upper internodes
and petioles. Stipules lanceolate, pale green, 15-27 x 6-10 mm, midrib prominent,
margin entire, apex narrowly pointed, caducous. Leaves alternate, 7(—11) cm apart;
petiole reddish, densely hirsute, 3—5 mm in upper leaves, 8—10 mm in lower leaves,
thickened at the base, slightly grooved above, in line with the midrib, /amina in juvenile
leaves silver-spotted between the veins, in mature leaves matt, plain light green with
a red patch at junction with petiole, beneath whitish or sometimes deep red or reddish
purple, oblanceolate, asymmetric, 12.5—17 x 8-10 cm, broad side 5—6.5 cm wide,
118 Gard. Bull. Singapore 55 (2003)
upper surface glabrous or with scattered dark red bristles between veins and on the
veins and midrib, the lower surface with lamina and veins glabrous or veins and
midrib minutely hispid, succulent in life, papery when dry, base cuneate and subequal
or unequal with the basal lobe of the larger side rounded, 0.5—1 cm long, margin
minutely toothed, apex acuminate; venation palmate-pinnate with 2 pairs at base and
3 lateral pairs along midrib, branching towards margin, impressed above, beneath
prominent and concolorous with lamina or sometimes red, basal lobe with 2 veins.
Plant protogynous. /nflorescences axillary, erect, cymose panicle, 10-16 cm long of
which peduncle is 0Q—3.5 cm long, greenish to dark red, minutely pilose, lowest node
with 1—2 female flowers, above up to 11 short branches 1—1.5 cm long with many
male flowers. Bracts green, ovate, 3-4 mm long. Male flowers with pale red pedicel
4—9 mm long, tepals 2, glabrous, deep pink or pale cream and deeper pink towards
the base, rotund, 6—7 x 6-7 mm or oval 5—8 x 3—7 mm, apex rounded; stamen cluster
3—5 x 2-3 mm with a stalk c. 1 mm long, stamens more than 35, filament c. 0.5 mm
long; anther pale yellow, obovoid, c. 1—-1.5 mm long, apex emarginate, dehiscing by
longitudinal slits in upper half. Female flowers with pedicel 7-9 mm long, reddish
green; ovary pale green sometimes with red veins on the wings, 13-18 mm long,
locules 3, each with 2 axile placentas, wings 3, equal, 3-5 mm wide; tepals 5, pale
peach to pale pink or green, sometimes deep ruby red at the base, slightly obovate,
6-12 x 3-8 mm, innermost smaller 4-10 x 3—7 mm, the margin entire or with 4-5
minute teeth towards the apex, apex acute, the outer surface sometimes with scattered
translucent hairs; styles 3, bifurcating, spirally twisted, yellowish-green to pale green,
2—6 mm long; stigma a glistening papillose spiral band. Fruits with stiff, decurved
pedicel 10-13 mm long, capsule glabrous, 2—2.7 x 1.8—2.8 cm, wings slightly unequal,
slightly pointed at tip, thinly fibrous, widest wing 8—11 mm wide, narrowest 6-8 mm
wide, dehiscing between locule and wing, style not persistent. Seeds barrel-shaped,
light brown, c. 0.4 mm long, collar cells c. 0.4 length of seed.
Distribution: Malaysia, Sarawak - endemic in Kuching limestone. Locally common
on G. Aup, G. Poing and G. Taba.
Habitat: At the base of limestone hills, in light shade.
Notes: Begonia lailana is named in honour of YABhg. Datuk Amar Puan Sri Dr Hajjah
Laila Taib, wife of the Chief Minister of Sarawak.
This species belongs to sect. Petermannia because the ovary has three locules
each with two placentas. It resembles B. oblongifolia in being a cane-begonia with
obovate leaves, paniculate male inflorescences and oblong fruits about 2 cm long.
However, it is distinct from this species in its leaves which are narrower, less than
twice as long as broad (they are c. 5 cm wide and more than twice as long as broad in
B. oblongifolia), in having fewer veins (8 in B. oblongifolia), larger, light red male
Begonias from Bau, Borneo 119
7mm
Figure 1. Begonia lailana Kiew and Geri.
A foliage; B partial inflorescence; C male bud; D male flower; E stamen cluster; F. stamen; G & H
female flower; I style and stigma; J. TS ovary.
120 Gard. Bull. Singapore 55 (2003)
flowers (3 mm long and green in B. oblongifolia) and wider wings (c. 5 mm wide in
B. oblongifolia).
Begonia lailana populations and individuals show some variation in leaf colour
and indumentum, and in ovary colour. Plants in some populations have completely
green leaves (e.g. at G. Kawa), in others most plants have green leaves with some
plants with red veins on the lower surface (e.g. at G. Tabai), or have most or some
plants with a deep red or purplish red undersides to the leaf (e.g. at G. Poing and G.
Aup, respectively). The shape of the silver spots on the juvenile leaves also varies
between populations: either discrete (e.g. at G. Kawa and G. Poing) or coalescent
(e.g. atG. Aup). Some populations have completely green ovaries (e.g. at G. Aup and
G. Poing), or the wings of the green ovary have fine red veins (e.g. at G. Kawa and G.
Tabai). Begonias are notoriously polymorphic for colour and patterning of the leaves
so these differences are not considered significant taxonomically.
While the stiff red bristles on the upper lamina surface are conspicuous, they
are not always present in all plants, e.g. the G. Aup population has glabrous leaves,
the Poing population has mostly glabrous individuals with a few with bristly leaves,
while the G. Kawa population has plants with bristly leaves. The female flower may
also vary in that the tepal margin may be entire or slightly toothed. However, this
difference does not correlate with any other character and is therefore not considered
taxonomically significant.
Other specimens examined: Gunung (G.) Aup Connie Geri et al. SBC 3756 (SBC),
Malcom et al. SBC 3303 (SBC); G. Batu Meekiong et al. SBC 2136 (SBC); G. Doya
Julia et al. SBC 2055 (SBC), Raymond et al. SBC 3451 (SBC), SBC 3473 (SBC); G.
Kawa Connie Geri et al. SBC 3753 (SBC), SBC 3592 (SBC, SING), Meekiong et al.
SBC 3114 (SBC); G. Lanyang Julia et al. SBC 2901 (SBC, SING); G. Podam Malcom
et al. SBC 3364 (SBC); G. Poing Julia et al. SBC 353 (SBC); Connie Geri et al. SBC
3755 (SBC), Malcom et al. SBC 1555 (SBC); G. Tabai Connie Geri et al. SBC 3757
(SBC), Julia et al. SBC 3414 (SBC).
4. Begonia pendula Ridl., J. Str. Br. Roy. As. Soc. 46 (1906) 257. Holotype:
Jambusan Ridley 11772 (K).
Sect. Petermannia
This species is endemic in the Kuching District where it is common on most limestone
hills. It creeps over vertical mossy cliff faces below the tree canopy with its thin stems
rooting at the nodes. Ridley (1906) named it ‘pendula’ describing its habit as having
trailing stems hanging down rock-faces. It is a decorative species as its bright green
leaves are variegated with silver-green elongated spots between the veins that in some
Begonias from Bau, Borneo 121
plants are discrete and in others coalescent. Some plants have leaves with the lower
surface maroon, others are green. The male flowers are deep pink and the female
flowers are red.
Distribution: Malaysia, Sarawak - endemic in Kuching limestone.
Other specimens examined: Bau Abang Mohrar et al. S52914 (SAR), Anderson 7799
(SAR, SING), Brooke 9891 (SING), Ridley 11774 (K); Bidi Clemens 20680 (SAR),
Yii et al. SSO381 (SAR); Bukit Boring Yii et al. S50360 (SAR); G. Apin Donny et al.
SBC 1360 (SBC), Jugah et al. S79954 (SAR), Makom et al. SBC 1107 (SBC), Malcom
etal. SBC 1107 (SBC); G. Aup Julia et al. SBC 568 (SBC), Malcom et al. SBC 3301
(SBC); G. Batu Meekiong et al. SBC 2126 (SBC), Meekiong et al. SBC 2126 (SBC),
SBC 2127 (SBC); G. Berloban Yii & Othman S$46234 (SAR); G. Doya Burtt BS1&7
(SAR), Julia et al. SBC 2027 (SBC), SBC 2144 (SBC), SBC 2154 (SBC), SBC 2162
(SBC), Sheavy et al. SBC 3056 (SBC); G. Jebong Donny et al. SBC 2095 (SBC),
Meekiong et al. SBC 2074 (SBC), Paul & Ilias $25623 (SAR); G. Juita Connie Geri
et al. SBC 3425, 3426 (SBC), Julia et al. 1293 (SBC); G. Pambur Julia et al. SBC
1263 (SBC) G. Podam Meekiong et al. SBC 3206 (SBC); G. Poing Julia et al. SBC
364 (SBC); G. Ropth Julia et al. SBC 2101 (SBC), Meekiong et al. SBC 3186 (SBC),
Shaevy et al. SBC 2426 (SBC); G. Selabor Sinclair SFN 38471 (SING); G. Setiak
Martin $38660 (SAR); G. Stulang Malcom et al. SBC 640 (SBC), SBC 643 (SBC);
G. Tabai Burtt BS158 (SAR), Julia et al. SBC 1293 (SBC), SBC 2513 (SBC), SBC
2563 (SBC); G. Tai Ton Julia et al. SBC 2677 (SBC), Stevens et al. 204A (SAR), Yii
et al. S51208 (SAR), $51210 (SAR); G. Tongga Donny et al. SBC 1777 (SBC); G.
Umbut Julia et al. SBC 2622 (SBC); Lobang Angin Yii et al. S51265 (SAR); Saburan
Anderson S$19119 (SAR).
5. Begonia rubida Ridl., J. Str. Br. Roy. As. Soc. 46 (1906) 256. Lectotype:
Jambusan Ridley 12393 (K, here designated).
Sect. Petermannia
As the name suggests, this species has ruby-red male flowers and its stems are also
bright red. It grows above the tree canopy on the shoulders of the summit, often on
peaty soil among sparse vegetation in light shade to full sunlight. Its leaves are thick
and succulent and are held vertically, unlike the leaves of the other cane-like begonias
that grow in shaded conditions, e.g. B. Jailana and B. congesta, the leaves of which
are held horizontally. This difference is probably a response to light conditions: the
horizontal leaf surface will intercept more light in shaded conditions, while leaves
held vertically will absorb less heat in bright sunlit conditions.
122 Gard. Bull. Singapore 55 (2003)
Distribution: Malaysia, Sarawak — endemic in Kuching limestone.
Other specimens examined: Bau Brooke 9832 (SING), Bidi Clemens 20681 (SAR);
Bukit Jebong Johnson S29403 (SAR), Paul & Ilias $25617 (SAR); Bukit Manok
Anderson $27416 (SING), Erwin & Paul S27416 (SAR); Bukit Numpang Chai &
Seng S22861 (SAR); Bukit Regu Paul et al. $37392 (SAR); G. Apin Donny et al.
SBC 1361 (SBC), Jugah et al. S79953 (SAR), Malcom et al. SBC 1108 (SBC); G.
Batu Meekiong SBC 2136, 2137 (SBC); G. Juita Connie Geri et al. SBC 3427 (SBC);
G. Meraja Julia et al. SBC 315 (SBC); G. Pambur Julia et al. SBC 1240 (SBC), SBC
1264 (SBC); G. Ropih Julia et al. SBC 2100 (SBC), Connie Geri et al. SBC 3516
(SBC); G. Setiak Martin $38670 (SAR), G. Taiton Julia et al. SBC 710 (SBC); G.
Tongga Denis et al. SBC 1722 (SBC), SBC 1732 (SBC); G. Umbut Julia et al. SBC
2584 (SBC).
6. Begonia speluncae Ridl., J. Str. Br. Roy. As. Soc. 46 (1906) 258. Lectotype:
Jambusan Ridley 11773 (K, here designated).
Sect. Reichenheimia
This begonia is, as its name suggests, a species that grows around cave mouths. It also
grows on dry rock cliff faces at the base of hills. It is endemic in the Kuching limestone
and has been collected from most hills in this area.
Doorenbos et al. (1998) placed this species doubtfully in sect. Reichenheimia
as the placenta type was not known. We confirm that it belongs to this section because
the ovary and fruit have three locules and the placenta is unbranched, 1.e. one per
locule.
Other specimens examined: Bidi Anderson 75 (K), Brooks s.n. (K) Clemens 20644
(K, SAR), Mjoberg 177 (K); Bukit Pait Erwin & Paul S27429 (SAR); Bukit Selabor
Paie S28044 (SAR, SING); Fairy Cave Rantai et al. S 70059 (SAR); G. Aup Julia et
al. SBC 707 (SBC); G. Bar Rantai et al. S66034 (SAR); G. Batu Meekiong et al. SBC
2135 (SBC); G. Doya Julia et al. SBC 2159 (SBC), Shaevy et al. SBC 3047 (SBC); G.
Jebong Lehman $30143 (SAR), Meekiong et al. SBC 2072 (SBC); G. Kawa Meekiong
et al. SBC 2797 (SBC); G. Lanyang Julia et al. SBC 1229 (SBC), SBC 2905 (SBC),
Denis et al. SBC 1439 (SBC); G. Manok Burtt BS132 (SAR); G. Meraja Julia et al.
SBC 301 (SBC); G. Podam Malcom et al. SBC 814 (SBC); G. Poing Julia et al. SBC
355 (SBC); Quop Hewitt s.n. (K); Sungai Serian Jacobs 5177 (SAR).
Begonias from Bau, Borneo 123
Acknowledgements
We are grateful to the ASEAN Regional Centre for Biodiversity Conservation for
funding The Biodiversity Survey of the Bau Limestone Project, of which this study is
a part; to the curators of the herbaria at Kew and the Forest Department, Kuching, for
permission to examine specimens in their care; to Sarawak Biodiversity Centre
colleagues, past and present, for their help in the field, to Raymond Sylvester Bohari
for preparing the illustration; and to M.J.S. Sands, Kew, for helpful suggestions on
improving the manuscript.
References
Doorenbos, J., M.S.M. Sosef and J.J.RE. de Wilde. 1998. The Sections of Begonia.
Wageningen Agriculture University Papers. 98-2.
Kiew, R. 1998. Niche partitioning in limestone begonias in Sabah, Borneo, including
two new species. Gardens Bulletin Singapore. 50: 161-169.
Pearce, K.G. 2003. Five new Begonia species (Begoniaceae) from the Niah National
Park, Sarawak, Malaysia. Gardens Bulletin Singapore. 55: 73-88.
Ridley, H.N. 1906. Begonias of Borneo. Journal of Straits Branch of Royal Asiatic
Society. 46: 246-261.
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Garden’s Bulletin Singapore 55 (2003) 125-162
New Species and New Records of Southeast Asian
Bulbophyllum (Orchidaceae)
J.J. VERMEULEN
Singapore Botanic Gardens
Singapore 259569
AND
F, 2, BYRNE
14 Clementi Cresent
Singapore 599517
Abstract
Seventeen new species of Bulbophyllum are described: B. tectipes (sect. Aphanobulbon, from
Sulawesi), B. pseudopelma and B. simplex (both sect. Desmosanthes, from Sulawesi), B.
flavorubellum (sect. Desmosanthes, from Sabah), B. decurrentilobum and B. iterans (both
sect. Hybochilus, from Sulawesi), B. furcillatum (sect. Hybochilus, from Sumatra), B.
anisopterum (sect. Monilibulbus, from Sabah), B. anakbaruppui and B. illecebrum (both sect.
Osyricera, from Sulawesi), B. uncinatum (sect. Polyblepharon, from Sulawesi), B. catillus, B.
lyriforme and B. sinapis (sect. Sestochilus, all from Papua New Guinea), B. coweniorum (sect.
Sestochilus, from Laos), B. incisilabrum (sect. Sestochilus, from Sulawesi), and B. turpis
(sect. Sestochilus, from Peninsular Malaysia). Bulbophyllum plumatum Ames, and B. spissum
J.J. Verm. are newly recorded for Peninsular Malaysia.
Introduction
The collection of orchid species in Singapore Botanic Gardens appears an inexhaustible
source of interesting plants. In this paper new species of the genus Bulbophyllum that
recently came to light are described. Several more new species of the same genus,
based on material from different sources (L, K) are added. Two Bulbophyllum species
that are newly recorded for Peninsular Malaysia are discussed.
126 Gard. Bull. Singapore 55 (2003)
Bulbophyllum sect. Aphanobulbon
Bulbophyllum tectipes J.J. Verm. & P. O’Byrne, sp. nov. — Fig. 1.
Bulbophyllum tectipes J.J. Verm. & P. O’Byrne, inter species rhizoma patenti ad
pendenti inflorescentia multiflora sect. Aphanobulbonis foliis 9.5—15 cm longis,
pedunculo omnino bracteis 3 vel 4 in duabus seriebus imbricatis acute carinatis
tecto notatum. — TYPE: Indonesia, Sulawesi, central part, SBG-O 1953 (SING,
holo.).
Roots creeping along the rhizome towards its point of attachment, then spreading
over the substrate. Rhizome patent to + pendulous with stiff, + patent branches, up to
80 cm long, 2—2.5 mm diam., sections between pseudobulbs 8-9 cm long, bracts
hardly persistent, distinctly keeled. Pseudobulbs distant, minute, cylindrical,
0.2—0.25 x 0.2-0.25 cm. Petiole 0.5—0.9 cm long. Leaf blade elliptic to ovate, 9.5—15
x 2.3—3.2 cm, index (length/width) 4.1—4.9; acute. /nflorescence patent, a rather dense
raceme, 4.4—10.4 cm long, 14—26-flowered. Peduncle 1.4—2.2 cm, bracts 3-4,
distichous, imbricate, the longest 14-20 mm long; acuminate, sharply keeled. Rhachis
3-8.2 cm. Floral bracts triangular, c. 3.8 x 1.5 mm, acute. Flowers not resupinate,
not fully opening, many simultaneously. Pedicel and ovary c. 2.2 mm long, basal
node + flush with the surface of the rhachis. Median sepal somewhat recurved, elliptic,
c.6 x 2 mm, index c. 3; acuminate, margins entire, base broadly attached; thin,
3-veined, glabrous. Lateral sepals recurved, free, oblique, triangular, c. 6 x 2.8 mm,
index 2.1—2.2; acute, otherwise as the median sepal. Petals porrect, elliptic, c.2.6 x 0.8 mm,
index 3.2—3.3; acute, margins entire, slightly papillose towards the tip, base rather
narrowly attached; thin but somewhat thickened towards the tip, 1-veined, surface
glabrous. Lip recurved + half-way, general outline + elliptic, c. 2.8 x 1.4 mm, index c.
2 (all without artificial spreading); acuminate, margins entire; rather thin, glabrous;
adaxially concave and with 2 conspicuous, rounded knobs near the base, with a median
furrow up to half-way the length of the lip, surface slightly convex towards the tip;
abaxially with a distinct, truncate ridge up to half-way the length of the lip. Column c.
1 mm long, column foot with a low ridge starting at the base of the stigma, and
continuing over 2/3 of the length of the column foot, column foot without a tooth just
above the ligament. Ste/idia porrect, triangular, c. 0.5 mm long, acute, along the lower
margin with a distinct, triangular, subacute tooth. Anther abaxially with a slight crest
distally, surface + glabrous, front margin not drawn out. Pollinia 2, semi-ellipsoid; no
appendages present.
Figure 1.Bulbophyllum tectipes J.J. Verm. & P. O’Byrne a. Habit; b. Flower; c. Flower analysis, from
left to right: median sepal, petal, lateral sepal, lip; d. Lip, left: adaxial side, right: abaxial side; e. Column
and lip, lateral view; f. Anther, above: abaxial side, below: adaxial side; g. Pollinia, left: single, right: a
pair. — All from SBG-O 1953 (living plant, spirit sample).
Southeast Asian Bulbophyllum 127
Figure 1. Bulbophyllum tectipes J.J. Verm. & P. O’Byrne
128 Gard. Bull. Singapore 55 (2003)
Colours: Plant glossy medium green. Flowers white.
Habitat: Epiphyte in montane forest, alt. 1 100—1300 m.
Distribution: INDONESIA: Sulawesi, central part (1 specimen seen).
Notes: Most similar to Bulbophyllum agapethoides Schltr. and B. falciculicorne
J.J. Sm., also from Sulawesi. The three species share a relatively inflexible rhizome.
B. tectipes is much larger vegetatively; the leaves are consistently almost twice as
long and more than twice as wide as in the species mentioned. Most characteristic,
however, are the conspicuous distichous, sharply keeled peduncle bracts that cover
most of the peduncle. B. rhizomatosum Ames & C. Schweinf. (Borneo, Peninsular
Malaysia), B. acuminatifolium J.J. Sm., and B. perpendiculare Schltr. (both from
Sulawesi) have limply pendulous rhizomes.
Bulbophyllum sect. Desmosanthes
Bulbophyllum flavorubellum J.J. Verm. & P. O’Byrne, sp. nov. — Fig. 2.
Bulbophyllum flavorubellum J.J. Verm. & P. O’Byrne, a Bulbophyllo flammulifero
rhachidi 0.5—1 cm longa floribus 4—8, stelidiorum margine basali sine ala antrorsa
differt. - TYPE: Malaysia, Sabah, Interior Province, Nabawan, Vermeulen &
Lamb 433 (K, holo., L, iso.).
Roots present along the entire rhizome. Rhizome creeping, 1—1.4 mm diam.,
sections between pseudobulbs 2—5 cm long, bracts hardly persistent. Pseudobulbs
distant, narrowly ellipsoid to subcylindrical, 1.7—3 x 0.3—0.6 cm. Petiole 0.3—0.4 cm
long. Leaf blade elliptic, 3.2-4.5 x 1.2—1.9 cm, index (length/width) 2.3—2.7; obtuse.
Inflorescences (sub-)erect, short racemes, 6—11 cm long, 4—8-flowered. Peduncle
5.5—10 cm, bracts 5—7, the longest 5—9 mm long. Rhachis 0.5—1 cm. Floral bracts
ovate, c. 4.8 x 2.2 mm, acuminate. F/owers not resupinate, not fully opening, several
open simultaneously. Pedicel and ovary c. 4 mm long, basal node + flush with the
surface of the rhachis. Median sepal somewhat recurved with an incurved top,
subtriangular, c. 9.5 x 2 mm, index 4.7-4.8; acute, margins entire, base broadly
attached; rather thin, 3-veined, glabrous. Lateral sepals recurved, free, oblique,
triangular, c. 9.5 x 2 mm, index 4.7-4.8; otherwise as the median sepal. Petals porrect,
ovate-triangular, c. 2.2 x 0.8 mm, index 2.7—2.8; obtuse, margins slightly erose distally,
base broadly attached; thin, 1-veined, glabrous. Lip recurved distally, general outline
+ ovate, c. 1.8 x 1.2 mm, index c. 1.5, acute, margins entire, very finely papillose
proximally; thick, surface glabrous; adaxially slightly concave near the base,
Figure 2. Bulbophyllum flavorubellum J.J. Verm. & P. O’ Byrne — a. Habit; b. Flower; c. Flower analysis,
from left to right: median sepal, petal, lateral sepal, lip; d. Lip, above: adaxial side, below: abaxial side;
e. Column and lip, lateral view; f. Anther, above: adaxial side, below: abaxial side; g. Pollinia, above:
two pairs, below: single pair. — All from Vermeulen & Lamb 433 (living plant, spirit sample).
Southeast Asian Bulbophyllum 129
Figure 2. Bulbophyllum flavorubellum J.J. Verm. & P. O’Byrne
130 Gard. Bull. Singapore 55 (2003)
with 2 obtuse ridges over most of the length of the lip, towards the base inconspicuous
and diverging, towards the tip distinct and fused to a V-shaped structure; abaxially
with a distinct, retuse ridge over most of the length of the lip, surface glabrous. Column
c. 2.2 mm long, stigma without a tooth at its base, column foot with a small, obtuse
knob just above the ligament. Stelidia porrect, falcate, triangular, c. 1 mm long, acute,
along the lower margin with an inconspicuous, deltoid, obtuse wing. Anther abaxially
without a crest, surface + glabrous, front margin drawn out into a flat, triangular,
acute tip. Pollinia 4, the inner slightly shorter than the outer, all obovoid, no appendages
present.
Colours: Sepals yellow proximally, bright orange red distally. Petals very pale
yellow. Lip orange. Column yellow. Flowers slightly scented.
Habitat: Terrestrial or as an epiphyte close to the forest floor, in lowland kerangas
forest. Alt. c. 300 m.
Distribution: MALAYSIA: Sabah (1 specimen seen).
Notes: On first sight, this could be taken for a exceptionally small form of
Bulbophyllum flammuliferum Ridl., from Peninsular Malaysia and Borneo. That
species, however, has a 2.5—10 cm long rhachis with 12-25 flowers, and a distinct,
antrorse, rounded wing along the lower margin of the stelidia.
Bulbophyllum pseudopelma J.J. Verm. & P. O’Byrne, sp. nov. — Fig. 3.
Bulbophyllum pseudopelma J.J. Verm. & P. O’Byrne, inter series speciebus
inflorescentiis racemosis Sect. Desmosanthes labello brevi lato, stelidiis
brevissimis notatum. — TYPE: Indonesia, Sulawesi, central part, SBG-O 3216
(SING, holo).
Roots present along the entire rhizome. Rhizome creeping or straggling away
from the substrate, 34.5 mm diam., sections between pseudobulbs 4.5—26 cm long,
bracts little persistent. Pseudobulbs distant, ovoid, 2.54 x 1.5—2 cm. Petiole 0.6—1.1 cm
long. Leaf blade elliptic-ovate, 9.5—17.5 x 1.9—2.5 cm, index (length/width) 5—7; obtuse.
Inflorescences many, often tufted, along the entire rhizome, (sub-)erect, lax racemes,
2-5.5 cm long, 2-4-flowered. Peduncle 1—1.8 cm, bracts c. 3, the longest 4-5.5 mm long.
Rhachis 0.4—3.5 cm. Floral bracts tubular proximally, 4-5 mm long, acute. Flowers
not resupinate, not fully opening, several open simultaneously. Pedicel and ovary c.
4.5 mm long, basal node + flush with the surface of the rhachis. Median sepal somewhat
recurved with an incurved top, elliptic-ovate, c. 7 x 3 mm, index c. 2.3—2.4; acuminate,
margins entire, base rather broadly attached; thin, 3-veined, glabrous. Lateral sepals
recurved, free, oblique, triangular, c. 7 x 2.5 mm, index c. 2.8; acute, base broadly
Figure 3. Bulbophyllum pseudopelma J.J. Verm. & P. O’ Byrne — a. Habit; b. Flower; c. Flower analysis,
from left to right: median sepal, petal, lateral sepal, lip; d. Lip, adaxial side; e. Lip, abaxial side; f.
Column and lip, lateral view; g. Anther, above: abaxial side, below: adaxial side; h. Pollinia, above: two
pairs, below: single pair. — All from SBG-O 32/6 (living plant, spirit sample).
Southeast Asian Bulbophyllum 131
Figure 3. Bulbophyllum pseudopelma J.J. Verm. & P. O’ Byrne
2 Gard. Bull. Singapore 55 (2003)
attached; otherwise as the median sepal. Petals porrect, ovate, c. 3.2 x 2.8 mm, index
1.1-1.2; rounded, margins entire, base rather narrowly attached; thin, 3-veined,
glabrous. Lip recurved distally, general outline ovate, c. 1.4 x 1.2 mm, index 1.1—1.2;
rounded, margins entire; thick and soft; adaxially slightly concave near the base, with
2 rather distinct, obtuse ridges proximally somewhat diverging and running parallel
up to c. 2/3 of the length of the lip, surface slightly convex towards the tip, glabrous;
abaxially with a distinct, retuse ridge over most of the length of the lip, surface finely
papillose in centre. Column c. 0.7 mm long, stigma with a transverse ridge at its base,
column foot with an obtuse knob just above the ligament. Ste/idia minute, porrect,
triangular, acute, along the lower margin with a distinct, antrorse, rounded wing.
Anther abaxially with a distinct crest, surface + glabrous, front margin not drawn out.
Pollinia 4, the inner slightly shorter than the outer, ellipsoid, the outer obovoid; no
appendages present.
Colours: Floral bracts greyish white. Flowers very pale greenish, pale yellow
towards the base.
Habitat: Epiphyte in montane forest, on the trunk and in the crown of a small
tree.
Distribution: INDONESIA: Sulawesi, central part (1 specimen seen).
Notes: Among the racemose species of sect. Desmosanthes, Bulbophyllum
pseudopelma is easily distinguished by the short and wide, fleshy lip and the very
short stelidia. In its general aspect this plant is reminiscent of B. reptans Lindl., a
species widespread in continental SE Asia. Apart from being sturdier, it is distinct in
the those characters mentioned above.
In the general aspect of its inflorescence, B. pseudopelma resembles some species
of sect. Pelma.
Bulbophyllum simplex J.J. Verm. & P. O’Byrne, sp. nov. — Fig. 4.
Bulbophyllum simplex J.J. Verm. & P. O’Byrne, a Bulbophyllo trifloro J.J. Sm. sepalis
9—10 mm longis sine apice incrassato differt. — TYPE: Indonesia, Sulawesi, central
part, SBG-O 3210 (SING, holo.).
Roots present along the entire rhizome. Rhizome creeping or straggling away
from the substrate, 0.8—1.5 mm diam., sections between pseudobulbs 5—28 cm long,
bracts moderately persistent. Pseudobulbs distant, ovoid, 1.5—2.4 x 0.6—0.9 cm. Petiole
0.1—0.3 cm long. Leaf blade elliptic-ovate, 3.5—6.5 x 1.1—1.5 cm, index (length/width)
3.1—5; obtuse. Inflorescences often along the entire rhizome, (sub-)erect, subumbellate
racemes, 1.8—2.5 cm long, 3—5-flowered. Peduncle 1.2—1.6 cm, bracts c. 2, the longest
Figure 4. Bulbophyllum simplex J.J. Verm. & P. O’ Byrne — a. Habit; b. Flower; c. Flower analysis, from
left to right: median sepal, petal, lateral sepal, lip; d. Lip, adaxial side; e. Lip, abaxial side; f. Column and
lip, lateral view; g. Anther, left: abaxial side, right: adaxial side; h. Pollinia, above: two pairs, below:
single pair. — All from SBG-O 32/0 (living plant, spirit sample).
Southeast Asian Bulbophyllum 133
Figure 4. Bulbophyllum simplex J.J. Verm. & P. O’Byrne
134 Gard. Bull. Singapore 55 (2003)
4—5 mm long. Rhachis 0.2-0.25 cm. Floral bracts triangular, 3.2—3.6 x 1.2 mm, acute.
Flowers partly resupinate, not fully opening, several open simultaneously. Pedicel
and ovary 2.6—4.2 mm long, basal node + flush with the surface of the rhachis. Median
sepal somewhat recurved with an incurved top, ovate, 9-10.5 x 3—3.3 mm, index 3-3.2;
acuminate, margins entire, base rather narrowly attached; thin, 5-veined, glabrous.
Lateral sepals recurved, free, oblique, triangular, 8—11.3 x 2.3-2.4 mm, index
3.44.4; acute-acuminate, base broadly attached; 3-4veined, otherwise as the median
sepal. Petals porrect, elliptic, 3.5—4 x 2—2.2 mm, index 1.7—1.9; acuminate, margins
slightly erose, base narrowly attached; thin, 3-veined, glabrous. Lip recurved, general
outline + oblong, 1.6—1.8 x 0.9—1.2 mm, index 1.5—1.8 (all without artificial spreading),
obtuse, margins finely papillose distally; rather thick; adaxially + flat near the base,
with 2 obtuse ridges over most of the length of the lip, towards the base inconspicuous
and diverging, towards the tip distinct and fused to a v-shaped structure, adaxial surface
finely papillose distally; abaxially with a distinct, retuse ridge over most of the length
of the lip, surface glabrous. Column c. 2 mm long, stigma without a tooth at its base,
column foot with a small, obtuse knob just above the ligament. Stelidia porrect, falcate,
triangular, c. 1 mm long, acute, along the lower margin with an inconspicuous, deltoid,
obtuse wing. Anther abaxially with a distinct crest, surface + glabrous, front margin
drawn out into a concave, rounded beak. Pollinia 4, the inner + as long as the outer,
all subtriangular, no appendages present.
Colours: Flowers completely white.
Habitat: Epiphyte in montane forest, in the crown of a small tree.
Distribution: INDONESIA: Sulawesi, central part (1 specimen seen).
Notes: Vegetatively and in the general aspect of the flowers Bulbophyllum simplex
is fairly similar to B. triflorum J.J. Sm., from Java. That species differs in having
consistently smaller flowers, with shorter sepals (3—4 mm long) with a thickened tip.
B. concinnum Hook.f., from Thailand and west Malesia, differs in the same characters,
as well as in having the pseudobulbs closer together and in having a narrower, ovate
lip.
Bulbophyllum sect. Hybochilus
Bulbophyllum decurrentilobum J.J. Verm. & P. O’Byrne, sp. nov. — Fig. 5.
Bulbophyllum decurrentilobum J.J. Verm. & P.O’ Byrne, a Bulbophyllo masarangico
Schltr. labello lobis lateralibus distaliter lobo mediali sine sinu transientibus
Figure 5. Bulbophyllum decurrentilobum J.J. Verm. & P. O’Byrne — a. Habit; b. Flower; c. Flower
analysis, from left to right: median sepal, petal, lateral sepal, lip; d. Lip, left: abaxial side, right: adaxial
side; e. Column and lip, lateral view; f. Anther, left: abaxial side, right: adaxial side; g. Pollinia, left:
single pair, right: two pairs. — All from SBG-O 3206 (spirit sample).
Southeast Asian Bulbophyllum 135
Figure 5. Bulbophyllum decurrentilobum J.J. Verm. & P. O’Byrne
136 Gard. Bull. Singapore 55 (2003)
differt. - TYPE: Indonesia, Sulawesi, central part, SBG-O 3206 (SING, holo.).
Roots mainly below the pseudobulbs. Rhizome creeping, 0.7—0.9 mm diam.,
sections between pseudobulbs 0.4—1 cm long, bracts not persistent. Pseudobulbs
distant, ovoid, 0.3—0.5 x 0.2-0.45 cm. Petiole up to 0.05 cm long. Leaf blade elliptic,
0.6—-1 x 0.2-0.25 cm, index (length/width) 3-4; acuminate. /nflorescence erect to
patent, 1.5—1.7 cm long, 1-flowered. Peduncle 0.8—0.9 cm, bracts 1, c. 1.5 mm long.
Floral bracts tubular proximally, c. 1.3 mm long, + acute. Flowers not fully opening.
Pedicel and ovary 1.1—1.2 mm long, basal node coinciding with the floral bract. Median
sepal recurved, ovate, c. 7 x 1.3 mm, index 5.3—5.4; long acuminate, margins entire,
base rather broadly attached; thin, 3-veined, glabrous. Lateral sepals oblique,
c.8x 1.3 mm, index c. 6.1—6.2; otherwise as the median sepal. Petals porrect, elliptic,
c. 1.8 x 0.7 mm, index c. 2.5—2.6; acuminate, margins erose distally, base rather
narrowly attached; thin, 1-veined, glabrous. Lip recurved in the distal half, general
outline ovate, c. 1.9 x 0.9 mm, index 2.1—2.2 (all without artificial spreading), margins
entire; glabrous, 3-lobed; midlobe ovate, rounded, rather thin; adaxially somewhat
convex and with a distinct, conical, obtuse knob near the base; abaxially without a
ridge; lateral lobes erect, attached along the proximal half, c. 0.5 mm wide atc. 1/3 of
the length of the lip, decurrent in front, rounded, thin. Column c. 0.6 mm long, stigma
without a ridge at its base, column foot without teeth. Ste/idia porrect, ovate, c. 0.3
mm long, + truncate. Anther abaxially with a distinct, finely papillose crest, surface
otherwise + glabrous, front margin drawn out into an incurved, triangular, subacute
beak with entire margins. Pollinia 4, the inner slightly more than half as long as the
outer, all obovoid; no appendages present.
Colours: Sepals and petals purple, whitish near the base. Lip purplish.
Habitat: Epiphyte in montane forest, on the trunk and on thin branches in the
crown of small trees. Alt. 1800—2500 m.
Distribution: INDONESIA: Sulawesi, central part (1 specimen seen).
Notes: Most similar to Bulbophyllum masarangicum Schltr., from the northern
part of Sulawesi (Minahassa). The main difference between the two is in the shape of
the lip: in B. decurrentilobum the front margin of the lateral lobes of the lip gradually
merges with the midlobe, in B. masarangicum it does so with a distinct sinus. B.
masarangicum also has smaller flowers, the sepals being 3—5 mm long.
Bulbophyllum furcillatum J.J. Verm. & P. O’Byrne, sp. nov. — Fig. 6.
Bulbophyllum furcillatum J.J. Verm. & P. O’Byrne, a Bulbophyllo camptochilo J.J.
Figure 6. Bulbophyllum furcillatum J.J. Verm. & P. O’Byrne — a. Habit; b. Flower; c. Flower analysis,
from left to right: median sepal, petal, lateral sepal, lip; d. Lip, above: adaxial side, below: abaxial side;
e. Column and lip, lateral view; f. Anther, left: adaxial side, right: abaxial side; g. Pollinia, above: two
pairs, below: single pair. — All from SBG-O 0922 (spirit sample).
Southeast Asian Bulbophyllum 137
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Figure 6. Bulbophyllum furcillatum J.J. Ve rm. & P. O’Byrne
138 Gard. Bull. Singapore 55 (2003)
Verm. sepalis lateralibus papillosis, labello longioribus angustioribusque differt.
— TYPE: Indonesia, Sumatra, Barisan Range, SBG-O 922 (SING, holo.).
Roots usually one below each pseudobulb. Rhizome creeping, 0.8—1 mm diam.,
sections between pseudobulbs 0.35—1 cm long, bracts not persistent. Pseudobulbs
distant, ovoid, 0.15—0.32 x 0.15—0.25 cm. Petiole up to 0.05—0.13 cm long. Leaf blade
elliptic to ovate, 0.6—1.2 x 0.4—0.7 cm, index (length/width) 1.5—2.3; mucronate.
Inflorescence erect to patent, 9-14 mm long, 1-flowered. Peduncle 0.3—0.7 cm, bracts
2, the longest 1.3—2.2 mm long. Floral bracts tubular proximally, c. 1.6 mm long,
acuminate. Flowers not fully opening. Pedicel and ovary 2.8-4 mm long, basal node
coinciding with the attachment of the floral bract. Median sepal + porrect, subtriangular,
c. 2.8 x 1.3 mm, index 2.1—2.2; acuminate, margins finely papillose distally, base
rather broadly attached; rather thin, thickened distally, 3-veined, adaxially finely
papillose distally. Lateral sepals slightly oblique, obovate, c. 2.9 x 1.8 mm, index c.
1.6—1.7; + acute, base broadly attached; adaxially finely papillose except the proximal
part of the upper half; otherwise as the median sepal. Petals porrect, + elliptic, c. 1.3
x 1 mm, index c. 1.3; subacute, margins erose distally, base narrowly attached; thin,
1-veined, glabrous. Lip recurved + half-way, general outline triangular, c. 1.5 x 0.5
mm, index c. 3; + obtuse, margins ciliate; thick; adaxially slightly concave near the
base and with a rounded, glabrous ridge that is bifurcate proximally and runs up to c.
1/3 of the length of the lip, with 2 more distinct, rounded, papillose ridges diverging
proximally and running up to c. 2/3 of the length of the lip, leaving a thin furrow in
between them; abaxially without a ridge, surface glabrous in the proximal third, with
long hairs elsewhere but with short hairs close to the tip. Column c. 0.9 mm long,
stigma with a distinct, conical, obtuse tooth at its base, column foot without teeth.
Stelidia porrect, triangular, c. 0.4 mm long, subacute, with a distinct, antrorse,
triangular, subacute tooth along the lower margin. Anther abaxially with a distinct
crest, surface + glabrous, front margin drawn out into a triangular, acute beak with
entire margins. Pollinia 4, the inner slightly more than half as long as the outer, narrowly
elliptic, the outer obovoid; no appendages present.
Colours: Sepals and petals yellowish green, but lateral sepals blackish purple in
the distal half except for the yellowish margins. Lip blackish purple.
Habitat: Understorey epiphyte on tree trunks and branches. Alt. 1400-1500 m.
Distribution: INDONESIA: Sumatra, Barisan Range (1 specimen seen).
Notes: Most similar to Bulbophyllum camptochilum J.J. Verm., from Brunei.
This species, however, has glabrous sepals, much narrower petals (index 2.4—2.5),
and a wider lip (index 1.3—1.4) that is subrectangular in general outline.
Figure 7. Bulbophyllum iterans J.J. Verm. & P. O’Byrne — a. Habit; b. top inflorescence; c. Flower; d.
Flower analysis, from left to right: median sepal, petal, lateral sepal, lip; e. Lip, left: adaxial side, right:
abaxial side; f. Column and lip, lateral view; g. Anther, left: abaxial side, right: adaxial side; h. Pollinia,
above: single pair, below: two pairs. — All from SBG-O 321] (spirit sample).
Southeast Asian Bulbophyllum 139
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Figure 7. Bulbophyllum iterans J.J. Verm. & P. O’Byrne
140 Gard. Bull. Singapore 55 (2003)
Bulbophyllum iterans J.J. Verm. & P. O’ Byrne, sp. nov. — Fig. 7.
Bulbophyllum iterans J.J. Verm. & P. O’ Byrne, a Bulbophyllo errato Ames floribus c.
11 mm longis, petalis adaxialiter distincte pubescentibus, labello longo gracili
differt. - TYPE: Indonesia, Sulawesi, central part, SBG-O 32/1] (SING, holo.).
Roots below the pseudobulbs. Rhizome creeping, 0.9—1.4 mm diam., sections
between pseudobulbs 0.4—0.9 cm long, bracts not persistent. Pseudobulbs close or
distant, ovoid, 0.4—0.9 x 0.3-0.8 cm. Petiole up to 0.06 cm long. Leaf blade elliptic to
ovate, 0.8—1.5 x 0.5—0.8 cm, index (length/width) 1.6—2; obtuse to acute. Inflorescence
a congested raceme, 3.54.3 cm long, 1—7-flowered. Peduncle erect to patent, 2.3-2.8 cm,
bracts 2—3, the longest 1.8—2.5 mm long. Rhachis slightly nodding, spindle-shaped,
up to 0.3 x 0.1 cm. Floral bracts triangular, 2.2—3 x 2-3 mm, acuminate. Flowers not
fully opening, each one developing only after anthesis of the previous. Pedicel and
ovary c. 2 mm long, basal node on a 1.5—2 mm long stump. Median sepal slightly
recurved, elliptic, 10.5—11.2 x 2.1 mm, index 5—5.4; acute, margins ciliolate, base
broadly attached; thin, 3—5-veined, glabrous. Lateral sepals subtriangular,
11-12.2 x 2.5—2.7 mm, index 4.44.6; otherwise as the median sepal. Petals porrect,
ovate, 7.6—7.9 x c. 1.9mm, index 44.2; acute, margins ciliate, base narrowly attached;
thin, 3-veined, adaxial surface with hairs except near the base. Lip slightly recurved
near the base, general outline ovate, slightly constricted near the base where the margins
approach each other over the adaxial surface, c. 5.8 x 1.2 mm, index 4.8-4.9; truncate,
margins ciliate; rather thick; adaxially concave near the base and with a distinct,
narrow, rounded, ridge running from the constricted part of the lip, near the base,
almost to the tip, adaxial surface glabrous; abaxially without a ridge, surface with
stiff hairs near the base, and with softer hairs up to + half way the length of the lip.
Column c. 1.8 mm long, stigma without a tooth at its base, column foot without teeth.
Stelidia porrect, triangular, c. 1 mm long, acute, with minute, antrorse, triangular,
obtuse tooth along the upper margin, close to the tip. Anther abaxially with a distinct,
papillose crest, surface otherwise finely papillose, front margin drawn out into an
incurved, truncate beak with entire margins. Pollinia 4, the inner c. 3/4 as long as the
outer, ovoid, the outer ellipsoid; no appendages present.
Colours: Flowers entirely dark purple.
Habitat: Epiphyte in montane forest, on the trunk and in the crown of small
trees. Alt. 1800—2500 m.
Distribution: INDONESIA: Sulawesi, central part (1 specimen seen).
Notes: While this species fits very well in sect. Hybochilus on account of the
floral characters it has a racemose inflorescence, a character that is very rare in that
Figure 8. Bulbophyllum anakbaruppui J.J. Verm. & P. O’ Byrne — a. Habit; b. Flower; c. Flower analysis,
from left to right: median sepal, petal, lateral sepal, lip; d. Lip, left: adaxial side, right: abaxial side; e.
Column and lip, lateral view; f. Anther, above: adaxial side, below: abaxial side; g. Pollinia, left: single
pair, right: two pairs. — All from SBG-O 3287 (spirit sample).
14]
Southeast Asian Bulbophyllum
Figure 8. Bulbophyllum anakbaruppui J.J. Verm. & P. O’Byrne
142 Gard. Bull. Singapore 55 (2003)
section and the assumedly related sect. Polyblepharon. Only two other species in
these sections display this character: Bulbophyllum erratum Ames (sect. Hybochilus,
from the Philippines) and B. vexillarium Rid. (sect. Polyblepharon, from Papua New
Guinea). B. iterans differs from both in having petals that are distinctly hairy on the
adaxial side, and the long, slender lip.
The name refers to the flowers that appear in succession on the inflorescence.
Bulbophyllum sect. Leptopus
Bulbophyllum spissum J.J. Verm.
Bulbophyllum spissum J.J. Verm., Blumea 41 (1996): 360. — TYPE: Indonesia, Lingga
Archipelago, Pulau Riau, O’Byrne LING 010 (L, holo.)
Habitat: Epiphyte in forest. Alt. 200-1000 m.
Distribution: MALAYSIA: Pahang, Pulau Tioman. INDONESIA: Lingga
Archipelago, Pulau Riau.
Notes: The Tioman plants represent the first record of this species for Malaysia.
Material seen: MALAYSIA. Pahang, Pulau Tioman, SBG-O 737 (SING), 739
(SING).
Bulbophyllum sect. Monilibulbus
Bulbophyllum anisopterum J.J. Verm. & P. O’Byrne, sp. nov.
Bulbophyllum anisopterum J.J. Verm. & P. O’Byrne, a Bulbophyllo stormii sepalo
medio sepalis lateralibus 1/4-plo longiore differt. - TYPE: Malaysia, Sabah,
Lamb 563/86 (K, holo.).
Bulbophyllum stormii sensu J.J. Vermeulen, Orch. Borneo 2 (1991) 227 (pro parte,
plate included), non J.J. Smith.
Roots below the pseudobulbs. Rhizome creeping, c. 1 mm diam., sections between
pseudobulbs c. 0.5 cm long, bracts not persistent. Pseudobulbs ovoid, basal half
prostrate on and fused to the rhizome so that the new pseudobulbs arise c. half-way
up the old, c. 0.9 x 0.4—0.5 cm. Petiole 0.5—0.8 mm long. Leaf blade elliptic, 1.8—2.5
x c. 0.5 cm, index (length/width) 3.5—5; obtuse. Inflorescence c. 6 cm long, 1-flowered.
Peduncle erect to patent, 3.5—-4 cm, bracts c. 3, the longest c. 2.5 mm long. Floral
bracts tubular, c. 2.5 mm, acute. Flowers not fully opening. Pedicel and ovary c. 18
mm long, basal node on ac. 2 mm long stump. Median sepal porrect, ovate, c. 9.5 x
2.8 mm, index c. 3.4; acuminate, margins entire, base rather widely attached; thin,
Southeast Asian Bulbophyllum 143
glabrous. Lateral sepals recurved, free, oblique, c. 32 x 6 mm, index 5.3—5.4; caudate;
otherwise as the median sepal. Petals porrect, ovate, c. 2 x 0.8 mm, index c. 2.5;
obtuse, margins entire, base widely attached; thin, glabrous. Lip slightly recurved c.
half-way, general outline ovate with a drawn-out top, c. 4.5 x 1.5 mm, index c. 3;
rounded, margins entire; rather thick, glabrous; adaxially concave towards the base,
convex and with the margins strongly recurved elsewhere; abaxially with a short,
retuse ridge near the base. Column c. 2 mm long, stigma with a slight callus at its
base, column foot without teeth. Ste/idia subulate, c. 1.5 mm long, acute. Anther
abaxially with a distinct, rounded, papillose crest, surface + glabrous, front margin
not drawn out. Pollinia 4, the inner less than half as long as the outer, all about ovoid,
without appendages.
Colours: Lateral sepals yellow, orange near the base. Petals translucent orange.
Lip purple red.
Habitat: Understorey epiphyte in wet montane forest, at 1200-1900 m alt.
Flowering observed in Jan, and Jun—Jul.
Distribution: MALAYSIA. Sabah, Ulu Padas (2 specimens seen).
Notes: In Vermeulen (1991) I thought this was an aberrant form of the widespread
and variable Bulbophyllum stormii J.J. Sm. Since then, I have found it again several
times, and it is now clear that it is a distinct species. It differs from B. stormii in
having a much shorter median sepal (about 1/3 to 1/4 of the length of the lateral
sepals. B. anisopterum is most similar to B. hodgsonii Henders., from Peninsular
Malaysia, but differs in having a much shorter median sepal, and caudate lateral sepals.
Bulbophyllum sect. Osyricera (= sect. Globiceps)
Bulbophyllum anakbaruppui J.J. Verm. & P. O’Byrne, sp. nov. — Fig. 8.
Bulbophyllum anakbaruppui J.J. Verm. & P. O’ Byrne, a Bulbophyllo hastifero Schltr.
pseudobulbis quadrangulatis, petalis latioribus, labello ovato differt.— TYPE:
Indonesia, Sulawesi, central part, SBG-O 3287 (SING, holo.).
Roots creeping along the rhizome towards its point of attachment, then spreading
over the substrate. Rhizome + pendulous with + patent branches, up to 80 cm long,
2.5—4 mm diam., sections between pseudobulbs 3.8—7.4 cm long, bracts moderately
persistent. Pseudobulbs distant, ovate, 2—3.4 x 0.9-1.3 cm, distinctly and obtusely
4-angled. Petiole 0.2-0.3 cm long. Leaf blade ovate, 8.5—12 x 2.1—3.1 cm, index
(length/width) 2.8—5.3; acute. Inflorescence porrect, a rather dense raceme, 3.8—6.2
cm long, 13—17-flowered. Peduncle 2—2.5 cm, bracts c. 4, the longest c. 6.5 mm long.
Rhachis, spindle-shaped, 2.2—3.2 x 0.38—0.43 cm. Floral bracts cordate, c. 4 x 5 mm,
acute. Flowers not resupinate, arranged in regular spirals, not fully opening, many
open simultaneously. Pedicel and ovary c. 2.2 mm long, basal node + flush with the
surface of the rhachis. Median sepal + porrect, obovate, c. 4.3 x 3.3 mm, index c. 1.3;
144 Gard. Bull. Singapore 55 (2003)
top cucullate, rounded, margins entire, ciliolate, base rather narrowly attached; rather
thick, 3-veined, surface very finely papillose adaxially towards the tip. Lateral sepals
adnate along the lower margins, oblique, ovate-triangular, c. 5.8 x 3.2 mm, index 1.8-1.9,
top flat, subacute, margins slightly papillose, base broadly attached; thick, otherwise
as the median sepal. Petals porrect, ovate, c. 3.4 x 1.7 mm, index c. 2; acute, margins
entire, ciliolate, base narrowly attached; rather thin, 1-veined, surface very finely
papillose. Lip slightly recurved + half-way, general outline ovate, c. 3 x 2.2 mm,
index 1.3—1.4; rounded, margins entire; rather thick, glabrous; adaxially slightly
concave near the base, with 2 inconspicuous, rounded ridges proximally somewhat
diverging and running from c. 1/3 of the length of the lip up to near the tip; abaxially
+ without a ridge near the base. Column c. 2.2 mm long, stigma without a tooth at its
base, column foot with 2 truncate wings near its base, and two more, retrorse, triangular,
obtuse ones just above the ligament. Stelidia minute, abruptly bent downwards but
with an antrorse. + acute tip, along the upper margin with a very large (far more
conspicuous than the stelidia themselves), antrorse, wedge-shaped wing with a broadly
rounded, erose top margin. Anther abaxially with a crest near its base, surface +
glabrous towards the base, coarsely papillose towards the tip, front margin drawn out
into a distinct, concave, triangular, rounded, coarsely papillose beak. Pollinia 4, the
inner + as long as the outer, all ellipsoid-ovoid; no appendages present.
Colours: Plant medium green. Flowers yellow, median sepal with thin red veins,
petals with thin red veins and margins, lip slightly suffused with red proximally.
Habitat: Epiphyte in a large tree in coffee and cocoa gardens near a village. Alt.
c. 1800 m.
Distribution: INDONESIA: Sulawesi, central part (1 specimen seen).
Notes: Suberect, patent or pendulous rhizomes occur only in two species of sect.
Osyricera: Bulbophyllum salaccense Rchb.f., from West Malesia, and B. hastiferum
Schltr, from Sulawesi. The latter is most similar to B. anakbaruppui, but differs in
having well rounded, not angular pseudobulbs, much narrower petals and a hastate
lip. In flower structure B. anakbaruppui resembles B. trigonobulbum J.J. Sm., from
Sulawesi. That species, however, has a creeping or straggling rhizome, with many
spreading roots. It also has narrower petals.
Named after the children of the Toraja village Baruppu, who collected the plant
from a large tree in the village gardens.
Bulbophyllum illecebrum J.J. Verm. & P. O’Byrne, sp. nov. — Fig. 9.
Bulbophyllum illecebrum J.J. Verm. & P. O’Byrne, a Bulbophyllo osyricera Schltr. et
Figure 9. Bulbophyllum illecebrum J.J. Verm. & P. O’Byrne — a. Habit; b. Flower; c. Flower analysis,
from left to right: median sepal, petal, lateral sepal, lip; d. Lip, left: adaxial side, right: abaxial side; e.
Column and lip, lateral view; f. Anther, left: adaxial side, below: adaxial side; g. Pollinia, left: two pairs,
right: single pair. — All from SBG-O 3215 (living plant, spirit sample).
Southeast Asian Bulbophyllum 145
Figure 9. Bulbophyllum illecebrum J.J. Verm. & P. O’Byrne
146 Gard. Bull. Singapore 55 (2003)
B. osyriceroidi J.J. Sm. sepalo mediali distincte concavo, petalis multo longioribus
angustioribusque, labello parte apicali spathulato differt. - TYPE: Indonesia,
Sulawesi, Central part, SBG-O 3215 (SING, holo.).
Roots mainly below the pseudobulbs. Rhizome creeping, 1.8—2 mm diam.,
sections between pseudobulbs 0.8—2 cm long, bracts barely persistent. Pseudobulbs
close together, ovate, 1—-3.5 x 0.7—1.1 cm, not angular. Petiole 0.7—2.8 cm long. Leaf
blade obovate-elliptic, 5.5-12 x 0.8-1.3 cm, index (length/width) 4.2—12.8; acute.
Inflorescence a rather dense raceme, c. 30 cm long, c. 24-flowered. Peduncle erect to
patent, c. 21 cm, bracts c. 5, the longest c. 9 mm long. Rhachis nodding at the base,
limply pendulous, not thickened, c. 9 cm long. Floral bracts + triangular, c. 3 x 3 mm,
acute. F/owers not resupinate, not fully opening, many open simultaneously. Pedicel
and ovary c. 3.2 mm long, basal node + flush with the surface of the rhachis. Median
sepal somewhat recurved, elliptic-obovate, c. 7 x 3.5 mm, index c. 2; top cucullate,
rounded, margins entire, papillose-ciliolate distally, base narrowly attached; very thin,
3-veined, surface glabrous. Lateral sepals adnate along the lower margins, recurved,
top part slightly incurved, oblique, ovate-triangular, c. 7.5 x 2.1 mm, index 3.5—3.6,
top flat, (sub-)acute, base broadly attached; thin, otherwise as the median sepal. Petals
recurved to spreading, ovate, c. 7 x 0.9 mm, index 12—13; acute, margins entire, slightly
erose near the tip, base narrowly attached; very thin, 1-veined, glabrous. Lip almost
straight, general outline subspathulate with a slightly widened base, c. 6 x 2 mm,
index c. 3, rounded, margins entire; proximally rather thick, distally very thick, almost
circular in section, glabrous; adaxially concave near the base, with a transverse wall
connecting the sides just above the ligament, top part distinctly convex; abaxially +
without a ridge near the base, top part distinctly convex. Column c. 2 mm long, stigma
without a tooth at its base, column foot with a distinct, transverse ridge near its base.
Stelidia minute, abruptly bent downwards but with an antrorse, + acute tip, along the
upper margin with a very large (far more conspicuous than the stelidia themselves),
antrorse + rectangular wing with a truncate, slightly erose top margin. Anther abaxially
with a crest near its base, surface + glabrous towards the base, papillose towards the
tip, front margin drawn out into a distinct, concave, triangular, obtuse, papillose beak.
Pollinia 4, the inner slightly shorter than the outer, ovoid; the outer ellipsoid; no
appendages present.
Colours: Sepals and petals translucent with thin, red veins. Lip greenish yellow,
margins red distally.
Habitat: Epiphyte in fairly mossy montane forest, on the trunk and in the crown
of small trees. Alt. 2300 m.
Distribution: INDONESIA: Sulawesi, central part (1 specimen seen).
Notes: Bulbophyllum osyricera Schltr., from Java and Sumatra, and B.
osyriceroides J.J. Sm., from Sumatra, are most similar to B. illecebrum. Both differ,
however, in having a more or less flat median sepal, much shorter (less than half as
Southeast Asian Bulbophyllum 147
long as the median sepal) and wider petals, as well as a lip with an ovate, rather than
a spathulate top part.
The diaphanous, widely spreading, antenna-like petals and the highly mobile,
very thick and brightly coloured lip suggest a baiting of pollinators by simulating
either prey or a sexual partner. The name refers to this.
Bulbophyllum sect. Polyblepharon
Bulbophyllum uncinatum J.J. Verm. & P. O’Byrne, sp. nov. — Fig. 10.
Bulbophyllum uncinatum J.J. Verm. & P. O’Byrne, a Bulbophyllo bisepalo et B.
stenochilo sepalis deorsum unciformibus, labello proximaliter gradatim dilatato
differt. — TYPE: Indonesia, Sulawesi, SBG-O 3835 (SING, holo.).
Roots all close to the base of the rhizome. Rhizome patent to hanging down
perpendicularly, up to 25 cm long, c. | mm diam., bracts not persistent. Pseudobulbs
ovoid-cylindrical, prostrate on and fused to the rhizome, so that the new pseudobulbs
arise from near the top of the old, 0.9-1.3 x 0.2—0.3 cm. Petiole up to 0.5 mm long.
Leaf blade ovate, 2—2.7 x 0.8—1.1 cm, index (length/width) 2.42.5; acute. Inflorescence
c. 0.6 cm long, 1-flowered. Peduncle porrect, c. 0.1 cm, bracts 1, c. 2 mm long. Floral
bracts tubular, c. 1.7 mm, acute. Flowers not fully opening. Pedicel and ovary c. 1.3 mm
long, basal node coinciding with the attachment of the floral bract. Median sepal
incurved at the top, ovate, c. 3.8 x 1.3 mm, index 2.9-3; subacute, margins entire,
ciliate towards the tip, base widely attached; thin, surface glabrous. Lateral sepals
recurved, fused along the lower margin, oblique, c. 3.3 x 1.2 mm, index 2.7—2.8;
obtuse; margins sparsely ciliate; otherwise as the median sepal. Petals porrect, ovate,
c. 1 x 0.7 mm, index 1.4—1.5; acuminate, margins entire, base widely attached; thin,
glabrous. Lip recurved at about 1/3 of its length, general outline ovate with a slightly
narrowed top, c. 2.3 x 0.6 mm, index 3.8—3.9; acute, margins ciliate towards the base,
ciliolate-papillose towards the tip; thin; adaxially slightly concave towards the base
and with a median ridge starting near the base and running over about 1/4 of the
length of the lip, surface convex and with the margins recurved towards the tip, entire
surface finely papillose; abaxially without a ridge near the base, glabrous. Column c.
0.9 mm long, stigma without basal tooth, column foot distinctly thickened and with a
retrorse, conical tooth near the base. Stelidia triangular, c. 0.7 mm long, acute, with a
slightly antrorse, obliquely deltoid, obtuse tooth along the lower margin. Anther
abaxially with a distinct, rounded, papillose crest, surface + glabrous, front margin
drawn out into a truncate beak with glabrous margins. Pollinia: 2, ovoid.
Colours: Sepals and petals yellowish near the base, purple towards the tip,
abaxially with minute, blackish spots. Petals translucent white. Lip purple.
Habitat: Found in coppiced roadside trees, at 500 m alt. Flowering observed in
148 Gard. Bull. Singapore 55 (2003)
Jan—Mar and Dec.
Distribution: INDONESIA. Sulawesi: central part (1 specimen seen).
Notes: Similar to Bulbophyllum stenochilum Schltr., from Papua New Guinea. B.
uncinatum differs in having only a low median ridge on the adaxial side of the lip,
without a retrorse tooth near the base of the lip.
Bulbophyllum sect. Sestochilus
Bulbophyllum catillus J.J. Verm. & P. O’ Byrne, sp. nov. — Fig. 11.
Bulbophyllum catillus J.J. Verm. & P. O’Byrne, a Bulbophyllo lyriforme et B. cheiri
columella lateraliter aspectu aliquot sigmatoide curvato differt. - TYPE: Papua
New Guinea, Jongejan, P. cult. 614 (L, holo).
Roots along the entire rhizome. Rhizome creeping, 4-5 mm diam., sections
between pseudobulbs 4—6 cm long, bracts with veins persistent as coarse fibres.
Pseudobulbs distant, ovoid, suberect, 1.5—2.0 x 0.8—1.2 cm. Petiole 1.8—2.5 cm long.
Leaf blade elliptic, 11-13.5 x 5.5—6.5 cm, index (length/width) 1.8—2.3; rounded.
Inflorescence c. 7 cm long, 1-flowered. Peduncle (sub-)erect, 1.2 cm, bracts c. 3, the
longest c. 12 mm long. Floral bracts tubular, c. 14 mm, cuspidate. Flowers not
resupinate, not fully opening. Pedicel and ovary 53—78 mm long, basal node on a
8—10 mm long stump, juga and valvae well-rounded. Median sepal somewhat incurved,
elliptic to obovate, 21—31 x 8—11.5 mm, index 2.2—2.9; acute to acuminate, margins
entire, base rather widely attached; rather thick, c. 1 1-veined, glabrous. Lateral sepals
with the lower margins running more or less parallel, free, oblique, subtriangular,
20-28 x 9-16 mm, index 1.7—2.2, base widely attached; otherwise as the median
sepal. Petals recurved to spreading, elliptic to obovate, 20—25 x 5.5—8 mm, index
3.2—3.8; (sub-)acute, margins entire, base rather narrowly attached; rather thin,
c. 7-veined, glabrous. Lip recurved near the base, general outline ovate with a drawn-
out top, 10-11 x 6-7 mm, index c. 1.4—1.9; obtuse, margins entire; top part thick,
elsewhere thin, surface glabrous; adaxially widely concave towards the base; abaxially
with a distinct, emarginate ridge over most of the length of the lip. Column 7-9 mm
long, slightly sigmoid in profile, stigma without basal tooth, column foot without a
tooth just above the ligament. Stelidia porrect, triangular, 1.5—2 mm long, obtuse,
without teeth along the lower margin. Anther abaxially with a very distinct, conical
crest, surface + glabrous, front margin drawn out into a rounded beak with erose
margins. Pollinia 4, the inner as long as the outer, all sub-triangular, without
appendages.
Figure 10. Bulbophyllum uncinatum J.J. Verm. & P. O’ Byrne — a. Habit; b. Flower; c. Flower analysis,
from left to right: median sepal, petal, lateral sepal, lip; d. Lip, left: adaxial side, right: abaxial side; e.
Column and lip, lateral view; f. Anther, left: abaxial side, right: adaxial side; g. Pollinia, left: a pair,
right: single. — All from SBG-O 3835 (spirit sample).
149
Southeast Asian Bulbophyllum
Figure 10. Bulbophyllum uncinatum J.J. Verm. & P. O’ Byrne
150 Gard. Bull. Singapore 55 (2003)
Colours: Unknown.
Habitat: Unknown.
Distribution: PAPUA NEW GUINEA (4 specimens seen).
Notes: The slightly sigmoid profile of the column is diagnostic within the group
of sect. Sestochilus with single, non-resupinate flowers. The lip widens abruptly
towards the base into two thin wings, as in Bulbophyllum lyriforme, and B. cheiri.
The top part, however, much shorter than in these species.
Bulbophyllum coweniorum J.J. Verm. & P. O’Byrne, sp. nov. — Fig. 12.
Bulbophyllum coweniorum J.J. Verm. & P. O’Byrne, a Bulbophyllo microglosso in
labelli latere adaxiali proximaliter pubescenti differt. -TY PE: Laos, SBG-O 3622
(SING, holo.).
Roots mainly below the pseudobulbs. Rhizome creeping, 5-6 mm diam., sections
between pseudobulbs 8—9 cm long, bracts with veins persistent as fibres. Pseudobulbs
distant, ovoid to ellipsoid, erect, 44.5 x 2.3—2.8 cm. Petiole 44.5 cm long. Leaf
blade elliptic, 9.5—19 x 4—5.7 cm, index (length/width) 2.3—3.4; obtuse. /nflorescence
9.5—11.5 cm long, 1-flowered. Peduncle erect to patent, 1.1—1.4 cm, bracts c. 2, the
longest c. 11 mm long. Floral bracts tubular, 15—18 mm, cuspidate. F/owers resupinate,
not fully opening. Pedicel and ovary 80-95 mm long, basal node on a 9-11 mm long
stump, juga and valvae well-rounded. Median sepal somewhat incurved, ovate, c.
23 x 9 mm, index 2.5—2.6; acute, margins entire, base widely attached; rather thin,
8—9-veined, glabrous. Lateral sepals recurved, lower margin without a fold, free,
oblique, ovate, c. 22 x 13 mm, index c. 1.7; acute-apiculate; otherwise as the median
sepal. Petals recurved to almost spreading, ovate, c. 18 x 6 mm, index c. 3; acute,
margins entire, base widely attached; rather thin, c. 5-veined, glabrous. Lip recurved
near the base, general outline + ovate, c. 8 x 8 mm, index c. 1; subacute, margins
entire; rather thin; adaxially concave towards the base, surface finely papillose-
pubescent towards the base, glabrous elsewhere; abaxially with a distinct, emarginate
ridge from base to tip, surface glabrous. Column c. 4 mm long, stigma without basal
tooth, column foot without a tooth just above the ligament. Stelidia minute, porrect,
deltoid, subacute, without teeth along the lower margin. Anther abaxially with a very
inconspicuous crest, surface + glabrous, front margin drawn out into a deltoid beak
with glabrous margins. Pollinia 4, the inner slightly shorter than the outer, all ovoid;
a large, fleshy lump of tissue attached to the inner pair.
Figure 11. Bulbophyllum catillus J.J. Verm. & P. O’ Byrne — a. Habit; b. Flower; c. Flower analysis, from
left to right: median sepal, petal, lateral sepal, lip; d. Lip, above: adaxial side, below: abaxial side; e.
Column and lip, lateral view; f. Anther, above: abaxial side, below: adaxial side; g. Pollinia, left: a single
pair, right: two pairs. — a. From LAE 77443 (herbarium material), with the flower reconstructed from
Jongejan cult. 1216; b. From Jongejan cult. 1216; c—g. From Jongejan cult. 614 (all spirit samples).
Southeast Asian Bulbophyllum tl
Figure 11. Bulbophyllum catillus J.J. Verm. & P. O’Byrne
152 Gard. Bull. Singapore 55 (2003)
Colours: Sepals and petals greenish with purple veins. Lip white, suffused with pinkish
purple.
Habitat: Unknown. Grows and flowers well in cultivation in lowland conditions
as well as at 1000 m alt. Flowering observed in Jan—Mar and Dec.
Distribution: LAOS: Locality unknown (1 specimen seen).
Notes: Most similar to Bulbophyllum microglossum Ridl. which, however, has a
glabrous lip. B. smitinandii differs in having two distinct patches of much longer
hairs on the adaxial side of the lip, as well as a distinct central callus.
Named after Bob and Chittima Cowen, Bangkok, Thailand, who successfully
flowered this species in their nursery.
Bulbophyllum incisilabrum J.J. Verm. & P. O’Byrne, sp. nov. — Fig. 13.
Bulbophyllum incisilabrum J.J. Verm. & P. O’Byrne, a Bulbophyllo rugoso Ridl. ovario
cristis non-sinuosis, sepalo mediali petalisque rotundato differt. — TYPE:
Indonesia, Sulawesi, central part, SBG-O 1879 (SING, holo.).
Roots mainly below the pseudobulbs. Rhizome creeping, 4.5—6 mm diam.,
sections between pseudobulbs 0.9-1.6 cm long, bracts with veins persistent as fine
fibres. Pseudobulbs close, ovoid-cylindrical, erect, 3.5—5.5 x 1.2—1.6 cm. Petiole
3-5.5 cm long. Leaf blade elliptic to obovate, 10—16.5 x 2.7-4.2 cm, index (length/
width) 3.2-4; obtuse. /nflorescence 11—12 cm long, 1-flowered. Peduncle erect to
patent, 2.6—3 cm, bracts c. 3, the longest 14-18 mm long. Floral bracts tubular, 15—18 mm,
cuspidate. Flowers resupinate, not fully opening. Pedicel and ovary 75—80 mm long,
basal node coinciding with the floral bract, juga and valvae sharply crested, crests
slightly curved towards the base of the flower. Median sepal somewhat recurved,
ovate, c. 20 x 12 mm, index 1.6—1.7; rounded, margins entire, base narrowly attached;
rather thick, 8—9-veined, glabrous. Lateral sepals + spreading, lower margin with a
distinct fold slightly below half-way the sepal, free, oblique, ovate-triangular, c. 23 x
12 mm, index c. 1.9—2; subacute, base broadly attached; otherwise as the median
sepal. Petals + recurved, ovate, c. 19 x 6 mm, index 3.1—3.2; rounded, margins entire,
base broadly attached; rather thick, 8—9-veined, glabrous. Lip sharply recurved near
the base and near the tip, general outline + ovate, c. 10.5 x 7.5 mm, index c. 1.4;
rounded, margins entire; thick, surface very coarsely warty near the tip; adaxially
concave near the base, with a median slit from base to tip, with 2 rather conspicuous,
rounded ridges diverging proximally and running up to c. 2/3 of the length of the lip,
surface glabrous; abaxially with a distinct, emarginate ridge from base to tip, surface
glabrous but with a finely papillose patch proximally along the median line. Co/umn
Figure 12. Bulbophyllum coweniorum J.J. Verm. & P. O’ Byme — a. Habit; b. Flower; c. Flower analysis,
from left to right: median sepal, petal, lateral sepal, lip; d. Lip, left: abaxial side, right: adaxial side; e.
Column and lip, lateral view; f. Anther, above: adaxial side, below: abaxial side; g. Pollinia, left: two
pairs, right: a single pair. — All from SBG-O 3622 (spirit sample).
Southeast Asian Bulbophyllum 153
~ooen & techy ah Bt
Figure 12. Bulbophyllum coweniorum J.J. Verm. & P. O’Byrne
154 Gard. Bull. Singapore 55 (2003)
c. 4mm long, stigma without basal tooth, column foot without a tooth just above the
ligament. Stelidia minute, porrect, triangular, acute, along the lower margin below
the base with a distinct, subtriangular, 3—3.5 mm long, obtuse tooth. Anther abaxially
with a rather distinct crest, surface + glabrous, front margin drawn out into a concave,
rounded beak with erose and papillose margins. Pollinia 4, the inner + as long as the
outer, all ovoid; no appendages present.
Colours: Sepals and petals yellow with red spots. Lip white, adaxially pink
towards the margins.
Habitat: Epiphyte in forest. Alt. 1100-1200 m. Flowering observed in Feb.
Distribution: INDONESIA: Sulawesi, central part (1 specimen seen).
Notes: This pretty species is most similar to Bulbophyllum rugosum Ridl. It
differs in having the crests on the ovary only slightly curved distally (not distinctly
sinuous), in having a rounded median sepal and petals, and in having widely spreading
lateral sepals.
Bulbophyllum lyriforme J.J. Verm. & P. O’Byrne, sp. nov. — Fig. 14.
Bulbophyllum lyriforme J.J. Verm. & P. O’Byrne, a Bulbophylo cheiri loborum
lateralium margine dorsali (proximali) concava labelli basi differt.— TYPE: Papua
New Guinea, Jongejan, P. cult. 282 (L, holo).
Vegetative parts not seen. Inflorescence c. 5 cm long, 1-flowered. Floral bracts
tubular, c. 6 mm, cuspidate. Flowers not resupinate, not fully opening. Pedicel and
ovary c. 40 mm long, basal node on a c. 5 mm long stump, juga and valvae well-
rounded. Median sepal somewhat incurved, subtriangular, 23—31 x 6-8 mm, index
3.84.5; acute, margins entire, base rather widely attached; rather thick, c. 9-veined,
glabrous. Lateral sepals recurved, free, oblique, 22—30 x 8-10 mm, index 2.5—3.4,
base widely attached; otherwise as the median sepal. Petals spreading to reflexed,
subtriangular, 21—28 x 2.24 mm, index 7—10.5; acute, margins entire, base widely
attached; rather thin, c. 5-veined, glabrous. Lip slightly recurved near the base, general
outline hastate-lyrate with a long drawn-out top part, margins slightly concave near
the base of the lip, 17-19 x 7-8 mm, index c. 2.2—2.4; obtuse, margins entire; drawn
out top part thick, elsewhere thin, surface glabrous; adaxially widely concave towards
the base; abaxially with a distinct, emarginate ridge over c. 1/5—1/4 of the length of
the lip. Column 6—8.5 mm long, stigma without basal tooth, column foot with a slight
callus just above the ligament. Stelidia porrect, triangular, 1—1.8 mm long, obtuse,
without teeth along the lower margin. Anther abaxially with a very distinct, flat crest,
Figure 13. Bulbophyllum incisilabrum J.J. Verm. & P.O’ Byrne —a. Habit; b. Flower; c. Flower analysis,
from left to right: median sepal, petal, lateral sepal, lip; d. Lip, left: adaxial side, right: abaxial side; e.
Column and lip, lateral view; f. Anther, left: adaxial side, right: abaxial side; g. Pollinia, left: single pair,
right: two pairs. — All from SBG-O /879 (living plant, spirit sample).
Southeast Asian Bulbophyllum 155
Figure 13. Bulbophyllum incisilabrum J.J. Verm. & P. O’ Byrne
156 Gard. Bull. Singapore 55 (2003)
surface + glabrous, front margin drawn out into a widely rounded beak with entire
margins. Pollinia 4, the inner as long as the outer, all about drop-shaped, without
appendages.
Colours: Unknown.
Habitat: Unknown.
Distribution: INDONESIA: Irian Jaya (4 specimens distributed from BO by J.J.
Smith, presumably from Irian Jaya). PAPUA NEW GUINEA (1 specimen seen).
Notes: Bulbophyllum lyriforme is most similar to B. cheiri Lindl.; it differs in
having the lateral lobes at the base of the lip slightly bent backwards, so that their
proximal margin is slightly concave. In B. cheiri this is invariably convex.
Bulbophyllum sinapis J.J. Verm. & P. O’ Byrne, sp. nov. — Fig. 15.
Bulbophyllum sinapis J.J. Verm. & P. O’Byrne, a Bulbophyllo macrantho in sepalis
lateralibus latioribus (indice 2—3.5 versus 1.5—1.7) differt. - TYPE: Papua New
Guinea, Jongejan, P. cult. 171 (L, holo).
Bulbophyllum aff. hahlianum Schitr.: O’ Byrne, Lowland Orchids Papua New Guinea
(1994) 456.
Roots along the entire rhizome. Rhizome creeping, 4-5 mm diam., sections
between pseudobulbs 8—12 cm long, bracts with veins persistent as coarse fibres.
Pseudobulbs distant, ovoid-cylindrical, suberect, 1.5—2.0 x 0.5—0.8 cm. Petiole 24.5
cm long. Leaf blade elliptic to obovate, 12—19.5 x 3.3—7.4 cm, index (length/width)
1.95.4; obtuse. [Inflorescence 5—12 cm long, 1-flowered. Peduncle (sub-)erect, 0.6—2 cm,
bracts 3—5, the longest 9-19 mm long. Floral bracts tubular, 10—16 mm, cuspidate.
Flowers not resupinate, not fully opening. Pedice/ and ovary 65—88 mm long, basal
node on a 13—16 mm long stump, juga and valvae well-rounded. Median sepal
somewhat incurved, elliptic to ovate, 26-28 x 8-12 mm, index 2.1—3.5; acuminate,
margins entire, base narrowly attached; rather thick, c. 11-veined, glabrous. Lateral
sepals with the lower margins running more or less parallel, often touching at the tip,
free, oblique, ovate-triangular, 16-22 x 10-13 mm, index 1.5—1.7, base widely attached;
otherwise as the median sepal. Petals recurved to spreading, elliptic to obovate,
18-30 x 4.8-8 mm, index 2.7—5; acute to acuminate, margins entire, base narrowly
attached; rather thin, c. 7-veined, glabrous. Lip recurved near the base, general outline
almost hastate, 5—6.5 x 5—6 mm, index c. 0.8—1.2; obtuse, margins entire; thick, surface
glabrous; adaxially hardly concave towards the base; abaxially with a distinct,
emarginate ridge from base to tip. Column 4.5—6 mm long, stigma without basal
Figure 14. Bulbophyllum lyriforme J.J. Verm. & P. O’ Byrne — a. Flower; b. Flower analysis, from left to
right: median sepal, petal, lateral sepal, lip; c. Lip, above: adaxial side, below: abaxial side; d. Column
and lip, lateral view; e. Anther, above: adaxial side, below: abaxial side; f. Pollinia, above: a single pair,
below: two pairs. — All from Jongejan cult. 282 (spirit sample).
Southeast Asian Bulbophyllum ‘57
~
shee
Figure 14. Bulbophyllum lyriforme J.J. Verm. & P. O’Byrne
158 Gard. Bull. Singapore 55 (2003)
tooth,column foot without a tooth just above the ligament. Stelidia porrect, triangular,
1—1.5 mm long, obtuse, without teeth along the lower margin. Anther abaxially with
a flat crest, surface + glabrous, front margin drawn out into a rectangular beak with
erose margins. Pollinia 4, the inner as long as the outer, all sub-triangular, without
appendages.
Colours: Flowers yellow.
Habitat: Epiphyte in lowland forest, coastal woodland. Alt. 0-100 m. Flowering
observed in Nov.
Distribution: PAPUA NEW GUINEA: New Britain (2 specimen seen).
Notes: Most similar to Bulbophyllum macranthum Lind1.; differs in having shorter
and wider lateral sepals (index 1.5—1.7; in B. macranthum 2-3.5). Bulbophyllum sinapis
has uniformly mustard-yellow flowers (hence the name), differing conspicuously from
the cream-coloured, heavily spotted flowers of B. macranthum.
Bulbophyllum turpis J.J. Verm. & P. O’ Byrne, sp. nov. — Fig. 16.
Bulbophyllum turpis J.J. Verm. & P. O’Byrne, a Bulbophyllo wrayi Hook. f. sepalo
mediali acuto, columna basi dente clitoriformi differt.— TYPE: Malaysia, Pahang,
Gunung Brinchang, SBG-O 060 (SING, holo.).
Roots mainly below the pseudobulbs. Rhizome creeping, 7-8 mm diam., sections
between pseudobulbs 2.8—3.5 cm long, bracts with veins persistent as fine fibres.
Pseudobulbs close or distant, ovoid, erect, 4-5 x 2.2—3 cm. Petiole 16—18 cm long.
Leaf blade elliptic, 28-29 x 4.8-—5 cm, index (length/width) 5.6—5.9; acute.
Inflorescence a rather dense raceme, 40-47 cm long, 40—50-flowered. Peduncle erect
to patent, 23—28 cm, bracts c. 7, the longest c. 28 mm long. Rhachis arching, 17-19
cm. Floral bracts triangular, 4.5—15 x 2—2.5 mm, subacute. Flowers + resupinate, +
secund, not fully opening, many open simultaneously. Pedicel and ovary 5—11 mm
long, basal node + flush with the surface of the rhachis. Median sepal + porrect,
ovate, c. 9.2 x 4.8 mm, index 1.9—2; acute, margins entire, base rather widely attached;
rather thick, 3-veined, glabrous. Lateral sepals somewhat recurved, oblique, triangular,
c. 9 x 5 mm, index c. 1.8, base rather narrowly attached; otherwise as the median
sepal. Petals porrect, subtriangular, c. 4 x 2 mm, index c. 2; obtuse, margins entire,
base broadly attached; thin, 1-veined, glabrous. Lip reflexed at c. 3/5 of its length,
general outline ovate with an oblong, drawn-out top part, c. 3 x 2 mm, index c. 1.5 (all
without artificial spreading), margins entire; glabrous, + 3-lobed; midlobe oblong,
truncate-emarginate, rather thick; adaxially concave and with 2 knob-shaped ridges
Figure 15. Bulbophyllum sinapis J.J. Verm. & P. O’ Byrne — a. Habit; b. Flower; c. Flower analysis, from
left to right: median sepal, petal, lateral sepal, lip; d. Lip, above: adaxial side, below: abaxial side; e.
Column and lip, lateral view; f. Anther, above: adaxial side, below: abaxial side; g. Pollinia, left: two
pairs, right: a single pairs. — All from Jongejan cult. 171 (spirit sample).
Southeast Asian Bulbophyllum 159
Figure 15. Bulbophyllum sinapis J.J. Verm. & P. O’Byrne
160 Gard. Bull. Singapore 55 (2003)
touching over a median furrow near the base, with 2 more rather inconspicuous, obtuse
ridges proximally somewhat diverging and running parallel from about 1/3 to 2/3 of
the length of the lip, the space in between channelled; abaxially with an inconspicuous,
retuse ridge up to about 1/3 of the length of the lip; /ateral lobes erect, attached along
the proximal third of the lip, semi-circular, c. 1 mm high, rounded, thin. Column c.
2.8 mm long, stigma with a transverse ridge at its base, a longitudinal median ridge
issuing from this, continuing up to + half-way the column foot, then splitting into two
spreading lobes ending in a retuse tooth just above the ligament. Stelidia porrect,
triangular, c. 1.2 mm long, obtuse to subacute, with a deltoid, acute tooth along the
upper margin and a similar, obtuse tooth along the lower. Anther abaxially with a
finely papillose crest, surface otherwise + glabrous, front margin drawn out into a
porrect, triangular, acuminate beak with papillose margins. Pollinia 4, the inner slightly
shorter than the outer, narrowly drop-shaped, the outer ovoid; no appendages present.
Colours: Rhizome scales densely spotted with dark red. Sepals yellow, stained
with red except near the base. Petals yellow. Lip yellowish green, top part yellow.
Flowers strongly scented, reminiscent of overripe apples.
Habitat: An understorey epiphyte in thick moss cushions in permanently wet
montane forest. Bulbophyllum tahanense Carr and B. evansii Henders., like similar
conditions and grow in the same area. Alt. 1600-2000 m. Flowering observed in Mar.
Distribution: MALAYSIA: Peninsula (1 specimen seen).
Notes: This species differs from Bulbophyllum wrayi Hook.f. in having acute
leaf tips, as well as a distinct bifurcate tooth on the column foot.
Bulbophyllum Incertae Sedis
Bulbophyllum plumatum Ames
Bulbophyllum plumatum Ames, Orch. 5 (1915) 184.— TYPE: Philippines, Mindanao,
Bukidnon Subprovince, BSC (L. Escritor) 21479 (not seen).
Bulbophyllum jacobsonii J.J. Sm., Blumea 1 (1935) 320.—TYPE: Indonesia, Sumatra,
West Coast, Soengai Dareh, Jacobson E.R. cult. 2202 (Theunissen) (not seen).
Habitat: In Malaysia found as an epiphyte in lowland swamp forest; in Sumatra
found at 1000 m altitude.
Distribution: MALAYSIA: Peninsula, Johor. INDONESIA: Sumatra.
PHILIPPINES: Mindanao.
Figure 16. Bulbophyllum turpis J.J. Verm. & P. O’ Byrne — a. Habit; b. Flower; c. Flower analysis, from
left to right: median sepal, petal, lateral sepal, lip; d. Lip, above: adaxial side, below: abaxial side; e.
Column and lip, lateral view; f. Anther, left: abaxial side, right: adaxial side; g. Pollinia, left: two pairs,
right: single pair. — All from SBG-O 0060 (living plant, spirit sample).
Southeast Asian Bulbophyllum 161
Simm
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Figure 16. Bulbophyllum turpis J.J. Verm. & P. O’Byrne
162 Gard. Bull. Singapore 55 (2003)
Notes: The Johor plants represent the first records of this species for Malaysia
and are the first fully documented records of this species in about 90 years. Plants of
Bulbophyllum plumatum probably originating from Indonesia appeared in trade in
some quantity a few years ago. The Johor plants differ slightly from these in having
the lateral sepals coarsely transversely wrinkled.
Material seen: MALAYSIA. Johor, Endau basin, km 56 on road to Mersing,
SBG-O 2533 (SING), Johor, Rompin area, Hon Quik Pin (WOC 17,2002) 01 (SING).
Garden’s Bulletin Singapore 55 (2003) 163-164
Re-examination of Vaccinium dialypetalum
(Ericaceae) — Erratum
S. P. VANDER KLOET
Department of Biology, Acadia University, Wolfville,
Nova Scotia, Canada B4P 2R6
In Vander Kloet (2002), I overlooked the fact that Sleumer had in 1941 already made
combinations for the taxa in question under Vaccinium. Therefore the names I proposed
are later homonyms and are illegitimate. The correct names for the taxa in question
should be:
Vaccinium ridleyi Sleumer, Bot. Jahrb. 71 (1941) 424. Agapetes parviflora Ridl., J.
As. Soc. Str. Br. 61 (1912) 26 non Dunn (1903), nec Vaccinium parviflorum Andr.
(1800); Agapetes micrantha Ridl., Fl. Mal. Pen. 2 (1923) 205, non Vaccinium
micranthum Dunal 1839; Vaccinium micranthum Vander Kloet 54 (2002) 181 non
Dunal (1839). Type: Ridley 15768 (holo SING!).
Synonym: Vaccinium pseudodialypetalum Ng, Gard. Bull. Sing. 28 (1976) 231. Type:
KEP 56673 (holo KEP).
Vaccinium urophyllum Merr., Pap. Mich. Ac. Sc. 19 (1934) 184. Type: Yates 2427
(holo MICH; iso NY, US).
Synonyms: Vaccinium longilingua Sleumer, Bot. Jahrb. 71 (1941) 424; Agapetes
perakensis Ridl., Fl. Mal. Pen. 2 (1923) 205, non Vaccinium perakense Ridl. (1918);
Vaccinium perakensis Vander Kloet, Gard. Bull. Sing. 54 (2002) 182 non. Ridl. (1918).
Type: Ridley 5532 (SING!).
Agapetes griffithii auct. non C.B.Clarke (1882): King, J. As. Soc. Beng. 74 (1906) II
ak
Acknowledgements
I acknowledge the help and advice of J.F. Veldkamp (L) and I.M. Turner (SING) in
the preparation of this erratum.
164 Gard. Bull. Singapore 55 (2003)
Reference
Kloet, S.P. Vander. 2002. Re-examination of Vaccinium dialypetalum (Ericaceae).
Gardens Bulletin Singapore. 54: 179-183.
Garden’s Bulletin Singapore 55 (2003) 165-171
OBITUARY
Mohamad Shah bin Hj Mohamad Noor
Mohamad Shah bin Hj Mohamad Noor, born in the staff quarters of the Singapore
Botanic Gardens, was the son of Hj Mohamed Nur bin Mohamed Ghous. Like his
father, he spent his working life in the herbarium exceeding his father’s record of 47
years’ service by one year.
Born on 22™ January 1935, he received his education at the Anglo-Chinese
School in Penang and the Tanjong Rhu Boys School in Singapore. After leaving school
in 1954, his father encouraged him to follow in his footsteps and so Mat Shah began
his career in the Herbarium in 1955 as a Plant Collector. When his father died in
1958, he took over his post as Herbarium and Museum Assistant. He rose to become
a Research Officer. No sooner did he retire in 1993 than he was re-employed until his
untimely death on 18" June 2003.
Mat Shah’s father was exceptionally diligent (Burkill, 1958) and never lost
the opportunity to collect plants. We even have a specimen of a banana he collected
from the market in Mecca while he was on the haj! Mohamed Nur undertook the
massive task of re-organising the herbarium to follow H.N. Ridley’s five-volume Flora
of the Malay Peninsula. Many of the covers in use today still bear his handwriting.
Besides his vast knowledge of plant names, he also amassed a vast collection of plants
from all over Peninsular Malaysia.
Mat Shah’s service in the Herbarium overlapped with his father’s for four
years, which gave him an invaluable grounding both in plant identification and
herbarium management. As a Plant Collector, Mat Shah accompanied the Director, J.
W. Purseglove, on three expeditions to Sarawak in 1955, 1956 and 1957. The longest
stint he spent in the field was when he joined the two Royal Society Expeditions to
Gunung Kinabalu in 1961 and 1964 (see Appendix). Except for these expeditions,
Mat Shah kept his own collecting books and numbering series and had collected
more than 5,000 specimens by the end of his career. Not only did he collect in Sabah
and Sarawak, he also collected extensively in the Peninsular Malaysia where he climbed
many of the major mountains (Gunung Belumut, G. Bujang Melaka, G. Lawit, G.
Ledang, G. Rabong, G. Tahan and G. Ulu Kali). He also collected continuously in
Singapore, from time to time being involved in extensive surveys, such as that of
plants in the Singapore Botanic Gardens in 1974 (96 numbers), the Seletar Reservoir
area in 1971 (51 nos.), MacRitchie Reservoir area in 1976 (77 nos.) and the Pierce
and Seletar Reservoir areas in 1981 (56 nos.).
He has recounted some of his experiences in the field (Waterson, 2003): >My
father taught me always to bring a mosquito net, because it can protect you against
centipedes and scorpions, even snakes. One night on Gunung Tahan about nine or ten
o’clock, I was lying under my net and I saw a very big cobra going up and over the
166 Gard. Bull. Singapore 55 (2003)
top of the net. One of the porters got a parang but I told him not to kill it. Because the
second thing my father taught me about the jungle was never to kill any animals. On
Gunung Rabung, there were tigers. We met a very big one, but the kampung people
assured us they were not man-eaters and never hurt anybody.”
Mat Shah worked through the most turbulent period in the herbarium’s history
as Singapore emerged from colonial status to independence and prosperity. There
were more changes in Director in his 48 years than in the previous 80 years. The nadir
was reached when rumours circulated worldwide that the Herbarium was no longer
valued and was up for sale!
He also experienced two major upheavals when the entire herbarium moved
(Kiew, 2003). The first in 1963 was forced because the herbarium building was
threatened with imminent collapse. The 400,000 specimens were moved inside the
cupboards, the Gardens’ labourers providing the muscle to move the heavy, solid teak
cupboards. The second was last year (2002) when the 650,000 specimens were moved
out so that the new state-of-the-art Herbarium could be built on the same site. Sadly,
Mat Shah will not be able to see the Herbarium settled into the new building in 2005.
For the second move, Mat Shah worked steadily for more than six months
with our team of ladies, Juriah Sabudin, Rohana Mohd Shari, Sariah Zelani and
Sugaranthara Davi, to bring the family and generic concepts up-to-date and number
the families according to D.J. Mabberley’s Plant Book and the genera and species
alphabetically. At the end of six months each box was numbered with its family and
sequence within the family. All the overflow of specimens (over 50,000) that had
accumulated over the years that could not be fitted into the old herbarium cupboards
was incorporated into the sequence. This systematic work paid off as it enabled the
10,000 boxes to be put straight onto the shelf in the correct order, all within 10 days.
This re-ordering the Herbarium is a lasting legacy to Mat Shah’s dedicated work.
Quiet, dedicated work characterised Mat Shah and his supervision of staff
over the years was firm yet harmonious. He will be remembered by the many visitors
to the Herbarium from all over the world for his courteous helpfulness. But it was
most particularly for his extensive knowledge of plant names (Kiew, 2001) that made
his service invaluable and led to his re-employment for eight years beyond retirement
age.
In 1993, he underwent a heart by-pass operation, which, although it curtailed
his plant collecting activities, did nothing to dampen his dedication to herbarium
work. He passed away peacefully in the early morning of 18" June 2003. He was due
to finally retire in August 2003.
His ability to identify plants and his knowledge of where plants grow in
Singapore will be sadly missed. In the words of Dr S.C. Chin, Director: “With his
passing, the tiny pool of specialist plant collectors and field botanists who can put a
name to most things green, shrinks further.”
He leaves his wife, Madam Jamilah binte Osman, whom he married in 1963,
three sons and a daughter, and four grandsons and three granddaughters.
167
Obituary-Mohd Shah b.Hj Mohd Noor
th
9.
)
H.M. Burkill, November 19
Mohamed Shah in the old Herbarium.
168 Gard. Bull. Singapore 55 (2003)
References
Burkill, H.M. 1958. Obituaries. Haji Mohamed Nur bin Mohamed Ghous, B.E.M.
Gardens Bulletin Singapore. 17: 331-336.
Kiew, R. 2001. What is this plant? The Herbarium’s role in plant identification.
Gardenwise. 17: 3-4.
Kiew, R. 2003. The Herbarium moves ... again! Gardenwise. 20: 13-15.
Waterston, R. 2003. Across the generations: Plant collectors and their careers at the
Botanic Gardens. Gardenwise. 21: 12, 14.
Ruth Kiew
Singapore Botanic Gardens
Singapore
Appendix: Mohamad Shah’s Collecting Expeditions Outside Singapore
(Bk Bukit, FR Forest Reserve, G Gunung, NP National Park, Sg Sungai)
1955
12-27 Sept. Sarawak — Kuching, Bk Kitang, Bau; Setapok, Sebanding; G Gading,
Lundu, Sematan, Kampung Pueh (with J.W. Purseglove, specimens numbered
under Purseglove’s collecting series)
1956
16 May-20 June. Sarawak — Bako NP, Telok Asam, Telok Tajor, Lokosan, Sg Serai,
Telok Wakang, Telok Delima, Telok Lakei, Telok Lomi, Bk Buan, Tau Range,
Bk Mayeng, Bk Merseng, Tatau (with Purseglove & H.B. Gilliland)
1957
2-18 Feb. Sarawak — Bako NP (with Purseglove)
6-9 May. Malacca — Fish Culture Research Station, Batu Berendam
18-22 Sept. Negeri Sembilan — Jendaram Estate, Nilai
1958
26 Jan—3 Feb. Pahang — Sabai Estate, Bentong
5-14 March. Perak — Bota Kiri FR, Ipoh (with Kadim bin Tassim)
2-9 April. Johore — Bk Paloh Estate (with Kadim)
Obituary-Mohd Shah b.Hj Mohd Noor 169
1959
1 Feb. Johore — Kota Tinggi Waterfall (with H.M. Burkill)
28 Feb-8 March. Kelantan —- Kampung Gobek, Tamangan (with Kadim)
24 Aug. Selangor/Pahang — Fraser’s Hill (with Burkill and Mohd Noor bin Jamaat)
21-28 Sept. Johore — Kuala Sedili, Teluk Jereh, Tanjung Lembu, Sg Sedili Besar
(with Mohd Noor and on 26" with Chang Kiaw Lan)
1960
15-19 Aug. Pahang — Fraser’s Hill (with Mohd Noor)
1961
18-26 May. Sabah — Pulau Gaya, Tanjung Aru, Bk Kepayang (with Kadim)
29May-16 Aug. Sabah — G Kinabalu (1* Royal Society Expedition, with Chew W.-
L, specimens numbered under the RSNB series)
1964
10 Jan-20 May. Sabah — G Kinabalu (2" Royal Society Expedition, with Chew)
1965
3-9 Dec. Perak — Maxwell Hill, Bk Berapit, Batu Hampar, Bk Kemunting (with Sidek
bin Kiah)
1968
16 Feb-7 March. Pahang — Taman Negara, Sg Tahan, Sg Teku, G Tuan Skeat, Kuala
Keniyan, Gua Luas, Sg Redap, Sg Belar (with herbarium of Forest Research
Institute Malaysia, KEP)
5 April. Johore — Mersing
1 Sept. Johore — Sg Pencharang, Bekok
1970
31 May. Johore — Sg Bantang, Labis F.R., Bekok (with E.A. Heaslett & Samsuri)
9-22 July. Pahang — Taman Negara, Ulu Sg Sat, Kuala Kelepah, Ulu Sg Sepia, Kuala
Aur, Ulu Sg Tembeling, Kuala Tahan (with KEP & Mohd Noor)
18-19 Aug. Johore — Bk Tangga Tujoh, Lak Heng Eststate, Kota Tinggi (with R. Hill,
Mohd Noor & Ahmad)
20-25 Sept. Johore — Sg Kahang, G Belumut (with KEP & Sanusi)
1971
29 Jan-4 Feb. Johore — Bk Pengantin, Sg Juasseh, Labis (with Ahmad)
22 April. Johore — Kota Tinggi Waterfall
5 Sept. Johore — Ulu Sg Segamat, Labis (with Samsuri Ahmad)
19 Sept. Johore — Padang Mulut, SE coast (with Ahmad)
170 Gard. Bull. Singapore 55 (2003)
1972
11-17 March. Kelantan — G Rabong, Ulu Sg Ketil (with KEP)
30 July. Johore — Sg Endau, G Janing (with Ahmad)
12-18 Aug. Pahang — Taman Negara, Bk Tersek, Kuala Trengan, Jeram Katak, Sg.
Keniyan, Genut Tabing (with Ahmad, on 18" with Miss Lee Wai Chin)
30 Aug-2 Sept. Pahang — Fraser’s Hill (with Miss Geh Siew Yin)
23 Nov. Johore — G. Lesong (with Ahmad)
1973
29 Jan-4 Feb. Kelantan — Gua Musang, Bk Brangkat, Bertam, G Brong, Kampong
Sta (with Mohd Ali)
11 Mar. Johore — G. Panti East (with Heaslett & Samsuri)
10-15 June. Selangor — Ulu Gombak, G. Ulu Kali, Anak Takun, G Bunga Buah
(with Mohd Ali)
23-30 Nov. Johore — G Lesong, Sg Rompin (with Ahmad)
1974
22-25 Feb. Kelantan — Bk Baka, Machang (with Ahmad)
27 Feb-1 March. Kelantan — Bk Yong, Pasir Puteh (with Ahmad)
31 May-4 June. Trengganu —G Tebu, Jabi, Sg Chabang Dua, Sg Belantan (with Ahmad
& Mahmud Awang)
1975
11-14 Feb. Perak — G Bujang Melaka, Sg Chenderiang (with Ahmad)
15 Feb. Pahang — Brinchang, Cameron Highlands (with Ahmad)
16-17 Feb. Perak — Sg Selung, Sg Hidup, Telok Anson (with Ahmad)
12-16 March. Trengganu — G Lawit, Kuala Sg Bok, Bk Yong (with British Museum
Gunung Lawit Expedition, with Ahmad and Samsuri)
11-17 July. Johore — G Ledang; Ma’okil, Cha’ah; Sg Batu Badak (with Samsuri)
23-28 Oct. Pahang — Bk Berkelah, G Serudom, Kuantan (with Samsuri & Sidek)
1976
3-5 March. Kelantan — Bk Bongkok, G Ayam (with Samsuri)
8 March. Kelantan — Ulu Sg Sat. (with Samsuri)
1978
9-13 June. Perak — G Bubu, Keledang Saiong FR (with Mahmud Awang)
5 Oct. Trengganu — Rantau Abang (with Samsuri)
22-26 Nov. Trengganu — Abang area, Jambu Bongkok, Gong Balai, Tanjung
Juara (with Sidek)
1981
15 Sept. Johore — Jason Bay, K. Sedili, Jemaluang Road (with Ali Rahim)
Obituary-Mohd Shah b.Hj Mohd Noor 171
1984
18-21 July. Trengganu — Bukit Tangga (with Mahmud)
1989
10-15 Feb. Sabah - G Kinabalu (to summit) and Poring (with E.P. Tay, specimens
numbered under Tay’s series)
20 Feb. Sabah — Nabawan (with E.P. Tay, specimens numbered under Tay’s series)
1991
23-26 July. Johore — Sg Linggui (with E.P. Tay and Ali Ibrahim, specimens numbered
under Tay’s series)
1993
2-5 May. Kedah — Sg Badak, Changlun, Bk Perangin; Bk Enggang, Sik.
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INSTRUCTIONS TO AUTHORS
Manuscripts: The Gardens’ Bulletin publishes original findings and reviews of progress in the
field of plant taxonomy, horticulture, and allied subjects. Contributions must be original and
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The Gardens’ Bulletin
Singapore
VOL. 55 (Part 2) December 2003 ISSN 0374-7859
CONTENTS
Kiew, R. and I.M. Turner
Pale Amy NMI SMe MMIC DO SUID ADIOLS Foo oo cic seen onete cenananiscnsemsumenecceel 1D
Hassan, I.
A note on Cordia premnifolia Ridl. (Boraginaceae) .................000000+-185
Mabberley, D.J.
New species of, and other notes on, Chisocheton and Walsura (Meliaceae)
cia oe ack ame RT shes Ds sv:anviees'esinpsasenesdenereeaetvabes 189
Gusman, G.
Arisaema fimbriatum (Araceae) and its intraspecific variation .............201
Middleton, D.J.
A revision of Dyera (Apocynaceae: Rauvolfioideae) ................2 606+. 209
Skornickova, J., M. Sabu and M.G. Prasanthkumar
Curcuma codonantha (Zingiberaceae) — a new species from the Andaman
DART NINA wine tira lias rir ties He i Call aig bes Bie. bh aii ane . Poaddanerrsa ses Shs 219
Zahid, M.S.
Three new species of Porterandia (Rubiaceae) from Mount Kinabalu,
eae Oh bed pes eked Die piesa od rretee sarc. tiwlsh dues. Vawenshenedssiove cheaters 229
Chung, R.C.K., E. Soepadmo and A.L. Lim.
The significance of pollen morphology in the taxonomy of Grewia and
Microcos (Tiliaceae) in Peninsular Malaysia and Borneo .....................239
Vermeulen, J.J. and P. O’Byrne
Six new species of Bulbophyllum (Orchidaceae) from Sulawesi ............257
Lee, S., A. Samsuri, P. Leong, Ali Ibrahim and A.T. Gwee
A botanical survey of Chek Jawa, Pulau Ubin, Singapore ......................271
Obituary
De Chane Kiaw Lan ...........:: 2 2e. sees oo. K.M Wong......309
Date of Publication: 15 December 2003
Published by
National Parks Board
Singapore Botanic Gardens
Cluny Road
Singapore 259569
Printed by
Print-Dynamics (S) Pte Ltd
Gardens’ Bulletin Singapore 55 (2003) 173-184
Are Any Plants Endemic to Singapore?
RUTH KIEW AND I. M. TURNER
Singapore Botanic Gardens, | Cluny Road, Singapore 259569
Abstract
Many taxa are reported in the literature as endemic to Singapore. Of the 19 investigated,
seven are shown to be endemic. Of these six, Bolbitis xsingaporensis (Lomariopsidaceae),
Flickingeria laciniosa (Orchidaceae), Spatholobus ridleyi (Leguminosae), Strychnos ridleyi
(Loganiaceae), Tectaria griffithii var. singaporeana (Dryopteridaceae) and Thunbergia
dasychlamys (Acanthaceae) are extinct, while Cryptocoryne xtimahensis (Araceae) is still
alive in the Bukit Timah Nature Reserve. These seven taxa represent a mere 0.3% of Singapore’s
vascular flora of about 2300 species. Anew combination is made for Hypobathrum coniferum
(Ridl.) Kiew (Rubiaceae).
Introduction
In Ridley’s 1900 enumeration of Singapore seed plants, the first listing for Singapore,
he noted 33 species as endemic to Singapore although he pointed out that most would
probably be discovered in neighbouring countries and this proved to be the case. In
addition, he indicated that some names he listed from the literature as coming from
Singapore were either invalid names or that Singapore had been wrongly given as the
place of origin. (The latter was not an unusual practice in the case of professional
plant collectors who did not want the true origin of a valuable plant to be known).
The preparation of the Flora of the Malay Peninsula, initiated at the Calcutta
Botanic Garden, but eventually driven by the indefatigable Ridley, both while he was
Director of the Botanic Gardens Singapore and later in retirement at the Royal Botanic
Gardens, Kew, resulted in the description of many new species, some of which were
known only from specimens collected from Singapore. The number of species named
in honour of Ridley and the number of rare species that he collected (see Appendix)
illustrate his diligence as a collector and that forest in Singapore was much more
extensive and less disturbed in his day
Since that period (1889-1925), a few new species apparently endemic to
Singapore have been described often based on very old collections. Significant among
these are two hybrids because hybrids often arise and survive in disturbed conditions
so they may be a reflection of disturbance in the remaining forested areas in Singapore.
174 Gard. Bull. Singapore 55 (2003)
Candidates for Endemic Species
Among the species listed by Ridley (1900), one has proved to be endemic to Singapore.
He described Dendrobium laciniosum, Orchidaceae (now known as Flickingeria
laciniosa (Ridl.) A.D.Hawkes) from two plants collected in 1891 from Pulau Seletar.
At one time, this species was confused with another orchid from Perlis, Peninsular
Malaysia, but Seidenfaden & Wood (1992) believe the Perlis plant to be different.
Incidentally, they erroneously record Pulau Seletar as in Johore, Peninsular Malaysia.
Later, Ridley described three more species and a variety as endemic to
Singapore: Didymocarpus perdita Ridl. (Gesneriaceae), Dischidia singaporensis Ridl.
(Asclepiadaceae), Petunga conifera Ridl. (Rubiaceae) and Spatholobus ferrugineus
(Zoll. & Moritzi) Benth. var. sericophyllus Ridl. (Leguminosae).
Didymocarpus perdita collected from Seletar in 1889, Ridley (1923) described
as ‘very rare, only 2 specimens seen in a ravine, now probably extinct’. This species
has proved to be the same as D. punticulata Rid|. (now known as Henckelia punticulata
(Ridl.) A. Weber), which grows in southern and eastern Peninsular Malaysia (Kiew,
1987).
Dischida singaporensis is similarly a very rare species known only from one
collection from the Changi Police Station in 1908. Later, D. singaporensis was collected
from Endau-Rompin, Johore (Kiew, 1987).
Petunga conifera, now known as Hypobathrum coniferum (Ridl.) Kiew (see
Appendix), in Ridley’s time was known from a single tree in the Botanic Gardens
Rain Forest, which at that time was called the Gardens Jungle (Ridley, 1923). Later,
a specimen of this species was collected from Johore, disproving its status as endemic
to Singapore. However, considering its extreme rarity and that is has not been
recollected for more than a hundred years, it can be concluded that the Singapore
population of this species is extinct.
Spatholobus ferrugineus var. sericophyllus was originally described from four
collections from Singapore (Ridley, 1922). However, this variety has since proved to
be widespread (Ridder-Numan & Wiriadinata, 1985). In Peninsular Malaysia, for
example, it has been collected from Perak, Selangor, Malacca and Negri Sembilan.
From publications from Calcutta, five new species were described that were
then known only from specimens collected in Singapore: Litsea ridleyi Gamble
(Lauraceae), Piper flavimarginatum C.DC. (Piperaceae), Polyosma ridleyi King,
Spatholobus ridleyi Prain ex King (Leguminosae) and Strychnos ridleyi King &
Gamble (Loganiaceae).
Litsea ridleyi, first collected in the 1880s, has been sporadically collected
since, the latest being in 1982 from the Bukit Timah Nature Reserve. However, several
specimens from Borneo (for example, Ridley s.n. 1892 from Bongaya and Senada
Brunei 54121 from Baram) are conspecific with this species, which is therefore not
Endemic Plants in Singapore i be,
endemic to Singapore.
Ridley (1924) reported the wild pepper, Piper flavimarginatum as only from
Singapore, where he described it as ‘very common but rarely flowering’. However, it
is widespread though not common in Peninsular Malaysia and has been collected
from as far afield as Langkawi, Perak, Pahang, Trengganu, Negri Sembilan and Johore.
It can therefore no longer be considered endemic to Singapore. It has not been
recollected in Singapore since 1933.
Another rare endemic, Polyosma ridleyi King (Escalloniaceae), was known
from a collection made by Ridley in 1892 from Seletar and another by Cantley in the
1880s without the locality specified. Much later it was shown by Saw (1989) to occur
in Johore and Brunei as well as Singapore.
Spatholobus ridleyi is a distinctive species. It has only been collected from
Singapore on three occasions and from nowhere else. It was last collected in 1899
and is an example of a rare Singapore endemic that is now extinct.
Strychnos ridleyi is known from just two specimens, both from Singapore,
the last collected in 1894. Ridley (1923) reported that it was ‘very rare in forests’.
The most recent revision for the region by Leenhouts (1962) confirmed it as a valid
species. S. ridleyi is another example of a rare Singapore endemic species that is now
extinct.
In the post-war period, new species were sporadically described either because
genera were being revised, such as the ferns Bolbitis singaporensis Holttum
(Lomariopsidaceae) and Tectaria griffithii (Baker) C.Chr. var. singaporeana Holttum
(Dryopteridaceae); Thunbergia dasychlamys Bremek. (Acanthaceae); Psychotria ‘sp.
9’ K.M.Wong (Rubiaceae) or, in one case, was recently discovered, Cryptocoryne
xtimahensis Bastmeijer (Araceae).
In 1947, Holttum described the fern Bolbitis singaporensis and noted that it
looked like a hybrid between what are now called Bolbitis appendiculata (Willd.)
K.Iwatsuki ssp. appendiculata and B. sinuata (C. Presl) Hennipman that all grew
together in one valley on Bukit Timah. In 1978, Hennipman formally designated this
fern as a hybrid, Bolbitis xsingaporensis Holttum. As it is only known from Singapore,
it is an endemic. Wee (1984) reported that it has not been seen ‘for a very long time’.
It was last collected in 1943. It is probably now extinct.
The case of the fern, Tectaria griffithii var. singaporeana is different. This
variety was only ever known from a single plant collected in 1908 from Bukit Timah.
The fact that it was not recollected by Holttum, who made an intensive study of ferns
in Singapore, nor by other botanists suggests that it was a single aberrant plant that
does not warrant taxonomic status. _
Thunbergia dasychlamys, a slender climber, is extremely rare and known
from only two specimens from Singapore, both collected by Ridley. It has not been
recollected since 1906. It is therefore a Singapore endemic, which is now extinct.
176 Gard. Bull. Singapore 55 (2003)
Psychotria malayana Jack is a common and widespread species with fruits
that dry conspicuously ribbed. However, some specimens that have fruits that are
almost smooth and leaves that are broader and the base not tapered into the petiole
were segregated from P. malayana by E.D. Merrill as recorded on his determination
slips dated 1950 on the specimens. These specimens were formally recognised as
distinct by Wong (1989) under the name Psychotria ‘sp. 9’. The status of this taxon
requires detailed investigation to decide whether Psychotria ‘sp.9’ is in fact a distinct
taxon or whether it is a southern form of P. malayana, something that is beyond the
scope of this study. It is not, however, endemic to Singapore as Merrill assigned a
specimen from Malacca (Ridley s.n. June 1892 from Gunung Mereng) to this taxon.
In Singapore, this taxon was collected from several localities giving the impression
that it was not rare. However, there are no specimens collected after 1906, which
strongly suggests that the Singapore population is now extinct.
Recently, Bastmeijer & Kiew (2001) described a new hybrid from Singapore,
Cryptocoryne xtimahensis Bastmeijer (Araceae) from the Bukit Timah Nature Reserve.
Its present status is endemic to Singapore as it has not been collected from elsewhere,
but considering how readily species of Cryptocoryne hybridise, it is likely that in
future it will be encountered in southern Peninsular Malaysia, where both the putative
parents grow.
Keng (1990) in his flora of Singapore cited two species, Polyosma kingiana
Schltr. (Escalloniaceae) and Sabia erratica P.M. v.d. Water (Sabiaceae) as endemic to
Singapore and both were listed as ‘strictly endemic to Singapore’ in Davis et al.
(1995). The inclusion of P. kingiana appears to be an error as this species, together
with the name it was originally described under, P. flavo-virens Ridl., was not recorded
from Singapore by Ridley (1922).
The case of Sabia erratica 1s an interesting one as van de Water (1980) clearly
stated in describing the species that the label on the type specimen at Kew, which
gave its origin as Bukit Timah, was clearly the wrong one as it described the plant as
a tree 30 m tall whereas Sabia is a genus of climbers. The data on the label recorded
the collector as Ngadiman (not Ngadiwan as given by van de Water) with the Field
Number 36149 (which is a Singapore Field Number, not a Kepong number as reported
by van de Water), collected from Tree No. 454 in Bukit Timah on 22 May 1940 and
the specimen was identified as Scaphium (Sterculiaceae). The collection belonging
to this label is in the Singapore Herbarium (SING) and represents a specimen of
Scaphium macropodum (Miq.) Beumee ex Heyne, which is indeed a tree. The type
specimen of Sabia was distributed from Kepong to Kew and at some point the label
from the Scaphium specimen was attached to the Sabia specimen. Van de Water
commemorated this mix-up in the specific epithet ‘erratica’ meaning wandering.
Since Sabia erratica was described, two other specimens have been identified
at Kew as belonging to this species. They are Makoto Togashi 622240 collected from
Endemic Plants in Singapore 177
Cameron Highlands, Pahang, at 6500 feet on | Feb 1962 and T & P 6/4 collected
from the Palas Tea Estate, Gunung Brinchang, Cameron Highlands, on | April 1979.
There is a duplicate of the latter specimen in SING. It is therefore now clear that
Sabia erratica is a very rare endemic from Cameron Highlands (probably confined to
Gunung Brinchang) and is not found in Singapore.
The status of Hanguana ‘Singapur’ (Hanguanaceae) suggested as a possible
new taxon (Tillich & Sill, 1999) is discussed in the Appendix. It is not endemic.
Discussion
Out of the potential candidates, only seven can be considered endemics (Table 1). As
mention above, two are hybrids that may have resulted from disturbance to the natural
habitats in the last hundred years. One variety, Tectaria griffithii var. singaporeana, is
doubtfully valid taxonomically as it is based on a single plant.
Singapore does not in fact possess a flora that is distinct from the surrounding
region but is a part of the flora found in southern Johore as well as showing affinities
with the northern flora of Borneo. The endemic status of the species in Table 1 may
not stand the test of time as the flora of Johore is relatively poorly collected compared
with that of Singapore and it is likely that these species also occur there.
It is interesting that three endemics are climbers, the life form most poorly
represented in herbaria, because of the inaccessibility of their foliage, flowers and
fruits high up in the canopy. Their endemic status in Singapore may therefore be an
artefact of more intensive collecting. (Singapore has always had more botanists per
km/? than any other country in the region). If a search is carried out in neighbouring
countries, these climbers can be expected to be found.
Table 1. Endemic species in Singapore
Species Life form No. localities & date Status
last collected
Bolbitis xsingaporensis fern 1 1943 extinct
Flickingeria laciniosa epiphyte | 1891 extinct
Spatholobus ridleyi climber Z 1899 extinct
Strychnos ridleyi climber d: 1894 extinct
Thunbergia dasychlamys _ climber 2 1906 extinct
Tectaria griffithii var. fern 1 1908 extinct
singaporeana
Cryptocoryne xtimahensis aquatic l extant
178 Gard. Bull. Singapore 55 (2003)
Most of these endemic species were always very rare and are known from
only one or two localities. Their extreme rarity meant that habitat destruction or
disturbance would tip them into extinction. Most of this extinction apparently occurred
about a hundred years ago as Singapore was intensively botanised in the 1920s and
1930s without these species being re-collected. Now only one of the seven still survives.
Seven endemics for a flora of about 2300 species of vascular plants represents
a very low level of endemism (0.3%), though it is higher (1.6%) if just ferns are
considered. One would not expect the Singapore flora to include a large number of
endemics as Singapore lacks many of the most biodiverse habitats with high endemism
of the region, such as mountains, limestone hills and ultramafic areas.
Conclusion
This survey shows that seven species are endemic to Singapore and that of these six,
Bolbitis xsingaporensis, Flickingeria laciniosa, Spatholobus ridleyi, Strychnos ridleyi,
Tectaria griffithii var. singaporeana and Thunbergia dasychlamys, are extinct; only
Cryptocoryne xtimahensis is still alive in the Bukit Timah Nature Reserve.
The other eight species thought at one time to be endemic have been shown,
as regional floras have become better known, to have a wider distribution and grow
outside Singapore. Seven of these were always very rare and have not been re-collected
in Singapore for many years, which indicates that their Singapore population is extinct,
although the species continues to survive elsewhere. These include Dischidia
singaporensis, Henckelia puncticulata, Hypobathrum coniferum, Piper
flavimarginatum, Polyosma ridleyi, Psychotria ‘sp. 9’ and Spatholobus ferrugineus
var. sericophyllus. A population of the eighth, Litsea ridleyi, still survives in Singapore
in the Bukit Timah Nature Reserve but is very rare (Maxwell, 1983).
The presence of the one extant endemic taxa and one rare species in the
Central Catchment Reserve reinforces its value as a protected conservation area for
the biodiversity of Singapore. The hybrid endemic to Singapore, Cryptocoryne
xtimahensis, grows in two adjacent small pools within the Bukit Timah Nature Reserve.
Its long-term survival is therefore vulnerable to any hydrological changes to that one
stream. Active monitoring and management, particularly in dry periods, is therefore
important.
Acknowledgements
This survey was prompted by Saifuddin Suran’s enquiry as to the status of the ‘endemic’
species mentioned in the Centres of Plant Diversity book. We thank Josef Bogner for
drawing our attention to the Hanguana ‘Singapur’; S. Chan for permission to work in
the Bukit Timah Nature Reserve; Saw L. G. for information on Sabia specimens at
Kepong; C. Chia and Saifuddin Suran, Biodiversity Centre, NParks, for searching for
Endemic Plants in Singapore 179
the location of Pulau Terijei; J.F. Veldkamp for help with tracing Hypobathrum names;
and the Keeper of Herbarium at Kew for permission for R. Kiew to study specimens
in his care, and to Lena Chan for comments on the manuscript.
References
Bastmeijer, J.D. and R. Kiew. 2001. A new Cryptocoryne hybrid (Araceae) from
Bukit Timah Nature Reserve, Singapore. Gardens’ Bulletin Singapore. 53: 9-
iF.
Davis, S.D., VLH. Heywood and A.C. Hamilton. 1995. Centres of Plant Diversity.
Vol. 2.Asia, Australasia and the Pacific. WWE and IUCN, U.K.
Keng, H. 1990. The Concise Flora of Singapore. Gymnosperms and Dicotyledons.
Singapore University Press, Singapore.
Kiew, R. 1987. The herbaceous flora of Ulu Endau, Johore-Pahang, Malaysia,
including taxonomic notes and descriptions of new species. Malayan Nature
Journal. 41: 201-234.
Leenhouts, P.W. 1962. Loganiaceae. Flora Malesiana. 1, 6: 293-387.
Maxwell, J.F. 1983. New and interesting plant records for Singapore. Gardens’
Bulletin Singapore. 35: 191-198.
Ridder-Numan, J.W.A. and H. Wiriadinata. 1985. A revision of the genus
Spatholobus (Leguminosae-Papilionoideae). Reinwardtia. 10: 139-205.
Ridley, H.N. 1900. The flora of Singapore. Journal of the Straits Branch of the
Royal Asiatic Society. 35: 27-196.
Ridley, H.N. 1922. Flora of the Malay Peninsula. Vol. 1. Reeve, U.K.
Ridley, H. N. 1923. Flora of the Malay Peninsula. Vol. 2. Reeve, U.K.
Ridley, H.N. 1924. Flora of the Malay Peninsula. Vol. 3. Reeve, U.K.
Rintz, R.E. 1980. The Peninsular Malaysian species of Dischidia (Asclepiadaceae).
Blumea. 26: 81-126.
180 Gard. Bull. Singapore 55 (2003)
Saw, L.G. 1989. Sabiaceae. Tree Flora of Malaya. 4: 426-431.
Seidenfaden, G. and J.J. Wood. 1992. The Orchids of Peninsular Malaysia and
Singapore. Olsen & Olsen, Denmark.
Tillich, H.-J. and E. Sill. 1999. Systematische Studien zur Morphologie und Anatomie
von Hanguana Blume (Hanguanaceae) und Flagellaria L. (Flagellariaceae), mit
der Beschreibung einer neuer Art, Hanguana bogneri spec. nov. Sendtnera. 6:
215-238.
Water, Th.P.M. van de. 1980. A revision of Sabia (Sabiaceae). Blumea. 26: 1-64.
Wee, Y.C. 1984. Common Ferns and Fern-Allies of Singapore. Malayan Nature Society
(Singapore Branch), Singapore.
Wong, K.M. 1989. Psychotria. Tree Flora of Malaya. 4: 396-399.
Appendix: Taxonomic notes with the inclusion of specimens in the Singapore
Herbarium (SING).
Bolbitis xsingaporensis Holttum
Basionym: Bolbitis singaporensis Holttum. Gard. Bull. Str. Settl. 11 (1947) 271.
Type: Bukit Timah — Hullett s.n. March 1882 (holo SING, iso SING).
Other specimens: Bukit Timah — C.G. Matthew 12567 1906 (SING), Holttum s.n. 27
Feb 1923 (SING), Holttum SFN 19799 12 Nov 1928 (SING), Holttum s.n. 12 Dec
1928 (SING), Holttum s.n. Feb 1943 (SING).
Cryptocoryne xtimahensis Bastmeijer. Gard. Bull. Singapore. 53 (2001) 11.
Type: Bukit Timah - Kiew et al. RK 5127 (holo SING).
Other specimen: Bukit Timah - P. Blanc 91-1 (SING).
Dischidia singaporensis Ridl. J. Str. Br. Roy. As. Soc. 61 (1912) 31.
Type: Changi Police Station — Ridley s.n. 1908 (lecto SING).
Note: The other specimen Ridley listed as belonging to this species, Rintz (1980)
identified as Hoya lacunosa Bl.
Flickingeria laciniosa (Ridl.) A.D.Hawkes.
Basionym: Dendrobium laciniosum Ridl. J. Linn. Soc. 32 (1896) 242.
Type: Pulau Seletar — Ridley s.n. 1891 (holo SING).
Endemic Plants in Singapore 181
Other specimen: Pulau Seletar — Goodenough s.n. 1891 (SING).
Hanguana ‘Singapur’
Josef Bogner (letter dated 5th Nov 2002) suggested that the Bukit Timah population
of Hanguana, based on his observations of a sterile plant in the Munich Botanic
Garden grown from seed collected in Bukit Timah, may be a new species, because
unlike H. malayana (Jack) Merr. (Hanguanaceae) with which it had previously been
identified, it was much smaller and did not produce long runners with young plants at
the tip. Tillich & Sill (1999) compared the morphology and anatomy of this sterile
plant, which they called H. ‘Singapur’, with three other Hanguana species from
Borneo, H. major Airy Shaw, H. bogneri Tillich & Sill and H. ‘Borneo’. However,
they did not compare the Bukit Timah plant with H. malayana.
Hanguana malayana can be a confusing species as its extreme forms look
very different. Plants that grow on lake or pond margins are robust plants that produce
long runners (the aquatic form), while plants that grow in forest in wet hollows or on
slopes (the terrestrial form) are more slender plants and their runners are much shorter.
However, since there are so many plants that are intermediate between these two
forms, Backer (1951) and Backer & Bakhuizen van den Brink (1968) concluded that
the aquatic and terrestrial forms belonged to a single variable species. Unfortunately,
neither of these characters (length of runners and robustness of the plant) can be seen
on herbarium specimens and data on labels is also usually lacking.
In deciding whether the Bukit Timah population represents a new species, it
was necessary to compare it with the complete range of variation in Hanguana
malayana from the aquatic to the terrestrial form. To this end, a study was initiated to
find, firstly, a suite of characters that can be examined from herbarium specimens,
that can be used to assess whether the terrestrial form is different from the aquatic one
in any characters other than robustness and runner length, and, secondly, to test whether
the Bukit Timah population, which grows in forest, falls within the range of variation
of either of these forms. This study was carried out by R. Kiew, A.T. Gwee, P.K.F.
Leong, Mohd Shah Mohd Noor and Samsuri Ahmad.
Based on the literature, eleven characters were selected that could possibly
separate the two forms, namely, leaf shape (ensiform v. lanceolate), indumentum (hairy
v. glabrous), petiole (present or absent), lamina length, width, and the ratio of width
to length, lamina tip (acute v. acuminate), inflorescence length (excluding stalk length),
length of lowest inflorescence branch, indumentum of inflorescence, fruit shape
(globose or fusiform) and fruit length. These were tested on herbarium specimens of
both forms from Peninsular Malaysia.
As with previous studies (Backer, 1951; Backer & Bakhuizen van den Brink,
1968), no character could be identified that would reliably separate the aquatic from
the terrestrial form. The characters either showed continuous variation, such as lamina
182 Gard. Bull. Singapore 55 (2003)
length and width or inflorescence length, or, if disjunct, such as fruit shape, did not
correlate with other characters.
The Bukit Timah Hanguana population grows in forest along streams and on
slopes, i.e. the habitat of the terrestrial form. Fertile specimens in the Bukit Timah
population were examined in the field for nine characters (leaf shape, leaf tip, petiole
and lamina length, lamina width, inflorescence length, length of the lowest
inflorescence branch, fruit shape and length). Results showed conclusively that they
fell within the range of the Peninsular Malaysian population although in lamina length,
they were in the lower range (mean length 57 cm, range 41-118 cm) compared with
mean length 76 cm, range 20.5—153 cm for H. malayana in Peninsular Malaysia.
Their smaller size might be explained by the nutrient-poor habitat of most of the
population, which grows on the coarse sand in an eroded streambed. Those on the
slope above the stream where there is a litter layer are larger plants but nevertheless
still fall within the range of variation seen in Peninsular Malaysia. There was no
difference for the other characters between the Bukit Timah and Peninsular Malaysian
populations. A few plants growing in water on the streambed had short runners up to
20 cm long.
This study therefore shows that the suggestion by Bogner that the Singapore
population of Hanguana represents a new species cannot be upheld as it falls within
the range of variation seen in H. malayana.
Backer, C.A. 1951. Hanguana. Flora Malesiana. Ser. 1. 4: 248-250.
Backer, CA. and R.C. Bakhuizen van den Brink. 1968. Hanguana. Flora of Java. 3:
20224:
Henckelia puncticulata (Ridl.) A. Weber.
Synonym: Didymocarpus perdita Ridl. J. Str. Br. Roy. As. Soc. 44 (1905) 54.
Type: Seletar — Ridley s.n. 3 Nov 1889. (holo SING).
Hypobathrum coniferum (Ridl.) Kiew, comb. nov.
Basionym: Petunga conifera Ridl. J. Str. Br. Roy. As. Soc. 79 (1918) 81; Ridl. FI.
Mal. Pen. 2 (1923) 85.
Synonym: Hypobathrum sp. 11 K.M.Wong. Tree Fl. Mal. 4 (1989) 355.
Invalid name: Hypobathrum coniferum Bakhf. in Keng, Concise Flora of Singapore
Gymnosperms and Dicotyledons (1990) 156.
Type: Singapore — Gardens Jungle Ridley 10722 1899 (holo SING).
Notes: Bakhuizen f. never published the combination cited by Keng, which is therefore
invalid. Hypobathrum coniferum resembles H. venulosum (Hook.f.) K.M.Wong but
differs in its very short inflorescences (up to 1.5 cm long) and its larger and broader
Endemic Plants in Singapore 183
leaves with more veins (15—18 x 6—7 cm with 7—9 veins) whereas H. venulosum has
long inflorescences up to 8 cm long and leaves 9-13 x 3.5—5 cm with 5-7 pairs of
veins. H. coniferum is therefore a valid species. All three specimens of H. coniferum
were collected in flower so its fruit remains unknown.
Other specimens: Singapore — Gardens Jungle Ridley 8427 April 1897 (SING);
Peninsular Malaysia — Johore, Sungai Kayu Kiah SFN 32366 10 March 1937 (SING).
Litsea ridleyi Gamble. Kew Bull. (1910) 317.
Type: Reservoir Wood - Ridley 5101 1893 (lecto SING (flowers), here designated).
Other specimens: Singapore (no locality) - Cantley s.n. 1880s (SING); Botanic
Gardens - Cantley 193 (SING); Gardens Jungle Hill HI038 11 Aug 1970 (SING);
Chan Chu Kang - Ridley s.n. 1894 (SING); Changi - Goodenough 3803 1892 (SING);
Bukit Mandai - Ridley 3836 1892 (SING), Ridley 4701 1892 (SING), Goodenough
5064 1893 (SING), Ridley10833 22 May 1900 (SING); Holland Road - Hullett s.n.
22 May 1893 (SING); Bukit Timah Nature Reserve - Hamzah Tambi H23 14 July
1971 (SING), Mohd Noor MN1878 22 April 1974 (SING), Shah & Samsuri 3946
1976 (SING), Maxwell 82-61 28 Feb 1982 (SING).
Notes: Among the Peninsular Malaysian species, it most resembles Litsea accedens
BI. (syn. L. singaporensis Gamble) in leaf and inflorescence characters but the latter
species is distinct in its fine reticulations that in dried leaves are raised.
The description in Ridley’s flora (1924) contains an error for petiole length,
which he describes as 0.15 inches long, when Gamble (1910) originally described it
as 1.5 cm long.
Piper flavimarginatum C.DC. Rec. Bot. Survey India. 4 (1912) 26.
Type: Chan-chu-kan Ridley 3772.
Other specimens: Bukit Timah — Ridley s.n. 1899 (SING), Corner SFN 3499 19 June
1938 (SING); Jurong — Corner s.n. 17 April 1933 (SING).
Polyosma ridleyi King. J. As. Soc. Beng. 66 (1897) 304.
Type: Seletar Ridley 3972 1892 (iso SING).
Note: Known from a single collection.
Psychotria ‘sp. 9’ K.M.Wong. Tree Fl. Mal. 4 (1989) 398.
Specimens: Singapore (without locality) - Cantley 2807 1880s (SING), Ridley 11272
1901 (SING); Chan Chu Kang - Hullett s.n. 1896 (SING); Bukit Mandai - Ridley
6556 1894 (SING); Bukit Panjang (?Puyan) Ridley 12530 1906 (SING); Bukit Timah - Ridley
11238 21 Oct 1900 (SING); Bukit Timah Road - Ridley 904 (SING); Jurong - Ridley
5422 March 1891 (SING).
184 Gard. Bull. Singapore 55 (2003)
Spatholobus ferrugineus (Zoll. & Moritzi) Benth. var. sericophyllus Ridl. Fl. Mal.
Peni4( 1922). 574.
Type: Upper Bukit Timah - Ridley 5578 (lecto SING).
Other specimens: Kranji - Ridley 6394 1894 (SING), Sinclair SFN 40687 27 July
1955 (SING); Bukit Mandai Goodenough s.n. 1890 (SING); Mandai Road - Ridley
s.n. 1900 (SING); Bukit Timah — Corner & Furtado s.n. 10 Nov 1929 (SING).
Spatholobus ridleyi Prain ex King. J. As. Soc. Beng. 66.II (1897) 80.
Type: Singapore Botanic Gardens Ridley 640] 1894 (iso SING).
Other specimens: Chan Chu Kang - Mat 6795 1894 (SING); Gardens Jungle - Ridley
s.n. 1899 (SING).
Strychnos ridleyi King & Gamble. J. As. Soc. Beng. 74.II (1908) 621.
Type: Tuas [Toas] Ridley 8313 4 May 1894 (isolecto SING).
Other specimen: Pulau ?Terijei - Goodenough s.n. 1891 (SING).
Notes: The name of the island (?Terijei) is illegible and we were unable to find an
island in the Singapore/Indonesian waters off Singapore that has a name that could
possibly match.
Tectaria griffithii (Baker) C.Chr. var. singaporeana Holttum. Blumea 35 (1990) 551.
Type: Bukit Timah - C.G. Matthew s.n. Jan 1908 (K).
Note: This variety is known from a single plant so there are no specimens at SING.
Thunbergia dasychlamys Bremek. Verh. K. Nederl. Akad. Wetensch. Afd. Natuurk.
Sect Tio 14 (1955) 53.
Type: Reservoir — Ridley s.n. 1906 (holo SING).
Other specimen: Tanglin - Ridley s.n. 1896 (SING).
Note: Bremekamp recorded ‘locality illegible’ for Ridley’s 1896 specimen. The label
in fact reads ‘hedges in Tanglin’.
Gardens’ Bulletin Singapore 55 (2003) 185-188
A Note on Cordia premnifolia Ridl. (Boraginaceae)
HASSAN IBRAHIM
Singapore Botanic Gardens
Singapore 259569
Abstract
Cordia premnifolia Ridl. is confirmed as a synonym of C. dichotoma G. Forst., not of C.
cochinchinensis Gagnep., which does not occur in Malesia. C. premnifolia is lectotypified.
Introduction
This paper arises out of the exercise to verify the status of type specimens in the
Singapore Herbarium (SING), in this case, to verify the type of Cordia premnifolia
Ridl. The family Boraginaceae was revised by Riedl (1997) for Flora Malesiana, in
which he regarded C. premnifolia as a synonym of C. cochinchinensis Gagnep. and
stated no types were known for C. premnifolia. By including C. premnifolia in the
latter, he considered that C. cochinchinensis occurs in Malesia. This paper discusses
whether C. premnifolia should be included in the more widely distributed C. dichotoma
G. Forst. as suggested by Ng (1989) or in C. cochinchinensis as suggested by Ried]
C9979.
Cordia premnifolia
The three specimens that Ridley (1915) cited when he described Cordia premnifolia
are all in the Singapore Herbarium. These specimens were collected in the Pahang
river region, Peninsular Malaysia.
Ridley (1915) described the leaves of C. premnifolia as ovate, obtuse or subacute,
6.5 x 4 in (16.5 x 10 cm), with the base as rounded or cordate and margins slightly
undulate to distantly serrulate. These characters distinguished it from C. griffithii
C.B.Clarke (now a synonym of C. dichotoma), which Ridley (1923) described as
having ovate-lanceolate acute to acuminate leaves with a slightly cuneate base,
1.5-3.5 x 1-1.75 in (3.8-9 x 2.5-4.4 cm).
A comparison of specimens of Cordia premnifolia and C. dichotoma from
186 Gard. Bull. Singapore 55 (2003)
Peninsular Malaysia and Singapore shows the three syntypes of the former to have
large, broad leaves up to 17 x 11 cm with a rounded to cordate base and entire to
slightly serrulate margins; while the majority of the latter has smaller, narrower leaves,
6.5—11 x 3.5—9 cm, with a cuneate base and wavy margin. However, some specimens
of C. dichotoma have broader leaves with a rounded base and an entire margin. In
addition, even on a single specimen of C. premnifolia there is a great range in leaf
size, for example, from 7.5—9.5 x 7 cm to 12.5—15 x 10-11 cm, thereby overlapping
with those of C. dichotoma. There are no differences in fruit shape and size and both
have the characteristic conical calyx cup. Taking the Peninsular Malaysian and
Singapore population of C. dichotoma as a whole, the C. premnifolia specimens,
although representing individuals with the largest leaves, fall within the range of
variation for that species. Ng (1989) was therefore correct in including C. premnifolia
as a synonym of C. dichotoma. He did not, however, lectotypify C. premnifolia.
Riedl (1997) made Cordia premnifolia a synonym of C. cochinchinensis
without giving a reason for his decision as well as without seeing any of the three
syntypes at SING and, because of this, he considered that C. cochinchinensis occurred
in Malesia.
Cordia cochinchinensis
In 1951, Johnston’s account of Cordia dichotoma highlighted its wide distribution
ranging from southern China and northern India, south through Indochina and Malesia
(including Peninsular Malaysia) to northeastern Australia. Gagnepain & Courchet
(1914) first described C. cochinchinensis from specimens collected in Cambodia,
Vietnam and Thailand (the Taleng Mountains in the Petchabouri region). Ried! (1997)
recorded C. cochinchinensis as occurring in Malesia presumably because he considered
C. premnifolia as a synonym of C. cochinchinensis. There is no other documentation
of this species being found elsewhere in Malesia. For example, it is not included in
the Flora of Java (Backer & Bakhuizen, 1965), the Tree Flora of Malaya (Ng, 1989),
the Tree Flora of Sabah and Sarawak (Dayang Awa, 1996) or Merrill’s 1923
Enumeration of Philippine Flowering Plants.
There are no specimens of Cordia cochinchinensis at SING but from the
descriptions by Gagnepain & Courcet (1914) and Johnson (1951), C. cochinchinensis
and C. dichotoma are very similar species with overlapping geographical ranges. The
main difference appears to be in the fruit: ovoid and up to 30 mm long and 22 mm
wide with a campanulate fruit calyx in C. cochinchinensis compared with ovoid and
about 22 mm long and 13 mm wide with a broadly cup-shaped to an inverted cone-
shaped fruit calyx about 6 mm long and 12 mm wide with obscurely lobed edges in
C. dichotoma (including C. premnifolia). Unfortunately, there is as yet no
Cordia premnifolia 187
revision of the Boraginaceae for the Flora of Thailand, where both species occur. It is
beyond the scope of this paper to conclude whether C. cochinchinensis and C.
dichotoma are conspecific. However, C. cochinchinensis does not occur in Peninsular
Malaysia and therefore not in Malesia either.
Cordia dichotoma G. Forst.
Cordia dichotoma G. Forst., Fl. Ins. Austr. (1786) 18, nr. 110; Johnston, J. Arnold
Arbor. 32 (1951) 8; Backer & Bakh. f., Fl. Java 2 (1965) 459; Ng, Tree Fl. Malaya 4
(1989) 60, Dayang Awa, Tree Fl. Sabah & Sarawak 2 (1996) 98, fig. 2; Riedl, Fl.
Malesiana I.13 (1997) 74.
Type: New Caledonia, Forster s.n., (not seen)
Synonym: Cordia premnifolia Ridl., J. Str. Br. Roy. As. Soc. 68 (1915) 12; Fl. Mal.
Pen. 2 (1923) 438.
Type: Peninsular Malaysia — Tanjong Antan, Pahang, Ridley 2158 (lecto SING, here
designated).
Other syntypes examined: Peninsular Malaysia - PAHANG: Pahang River Burn
Murdoch 186 (SING), Kuala Lipis Machado 11567 (SING).
Notes: Ridley 2158 is chosen as the lectotype among the specimens it is the most
complete having mature fruits.
Acknowledgements
I thank the Asean Regional Centre for Biodiversity Conservation for providing the
grant for the Singapore Type Project and Ruth Kiew, Singapore Botanic Gardens, for
support and guidance in preparing this paper.
References
Backer, C.A. and R.C. Bakhuizen van den Brink 1965. Boraginaceae. Flora of Java.
2: 457-459.
Dayang Awa, A.L. 1996. Boraginaceae. Tree Flora of Sabah and Sarawak. 2: 97-100.
Gagnepain F. and L.D.J. Courchet. 1914. Boraginacées. Flore Generale |’ Indo-Chine.
4: 203-204.
188 Gard. Bull. Singapore 55 (2003)
Johnston, I.M. 1951. Studies in the Boraginaceae, XX, Representatives of three
subfamilies in Eastern Asia. Journal of the Arnold Arboretum. 32: 8-11.
Merrill, E.D. 1923. Boraginaceae. An Enumeration of Philippine Flowering Plants.
3: 373-374.
Ng, F.S.P. 1989. Boraginaceae. Tree Flora of Malaya. 4: 60-62.
Riedl, H. 1997. Boraginaceae. Flora Malesiana. I, 13: 68-79.
Ridley, H.N. 1915. New or Rare Malayan Plants. Journal Straits Branch of Royal
Asiatic Society. 68: 12.
Ridley, H.N. 1923. Boragineae. Flora of the Malay Peninsula. 2: 438-439.
Gardens’ Bulletin Singapore 55 (2003) 189-200
New Species of, and Other Notes on, Chisocheton and
Walsura (Meliaceae)
D. J. MABBERLEY
Nationaal Herbarium Nederland, University of Leiden, The Netherlands;
'National Herbarium of New South Wales, Royal Botanic Gardens Sydney, Mrs
Macquaries Road, Sydney 2000, Australia'
Abstract
Chisocheton maxilla-pisticis Mabb., from Sabah, Malaysia, and Palawan, Philippines, and C.
velutinus Mabb., from Sarawak, Malaysia, and Brunei, are described as new. A third new
Chisocheton species, from Sarawak, is so far known only from a single fruiting specimen.
Walsura pinnata Hassk. sensu T.Clark includes a number of distinctive Bornean entities, two
of which are here afforded or re-afforded specific rank: Walsura decipiens Mabb., from north-
ern Borneo and W. grandifolia Ridl. from the Sarawak limestone. Chisocheton nicobarianus
Debnath & Sreek. is reduced to C. macrophyllus King, which is lectotypified here, while
Walsura yunnanensis C.Y.Wu from China is reduced to W. pinnata sensu lato (W.
cochinchinensis (Baill.) Harms).
Introduction
In preparing the text of Meliaceae for the Tree Flora of Sabah and Sarawak, it be-
came clear that new Chisocheton species signaled in the author’s monograph (1979;
see also 1995) could now be described using materials collected since its publication.
Examination of Borneo materials assigned by Clark (1994) to Walsura pinnata Hassk.,
a perplexingly variable species, has shown that two distinct species, one of them new,
are better segregated from it. In passing, notes on allied taxa from outside Borneo are
included.
Chisocheton Blume
1. Chisocheton maxilla-pisticis Mabb., sp. nov., Fig. |
Chisocheton, C. macrophyllo King affinis, indumento dense-fulvo, foliolis costis
paucioribus, pseudogemmula hiante vix circinnata, maxilla pisticis simulante, floribus
pusillioribus, differt. Typus: Malaysia, Sabah, Telupid, Pinangah F.R., 10 July 1993,
Sawan Tingki SAN 136830 (holo SAN!, iso KEP!, SAR!).
' Address for correspondence
190 Gard. Bull. Singapore 55 (2003)
Rather pachycaul tree with sparsely branched crown to 45 m tall; fluted bole to 90 cm
diameter with steep buttresses to 1(—3) m tall. Bark smooth to scaly, markedly
lenticellate; inner bark reddish brown; sapwood pale yellowish white. Twigs brown,
with longitudinal cracks, cicatrices conspicuous, latex white. Leafy twigs 1.2-1.5 cm
diameter, fulvous-tomentellous. Leaves in dense terminal spirals, to 1.3 m long,
pseudogemmulate; petioles 8.5—16.5 cm long, more or less angled or grooved, like
rachis, indumentum sericeous to pilose; leaflets in up to at least 17 pairs, oblong-
ovate, weakly falcate, to 21 x 8.5 cm, rather fleshy (Pennington), young ones brownish
or reddish, densely pilose on midrib and costae, also abaxially scattered between
veins, bases asymmetrical, rounded or, particularly in juveniles, cuneate, apices acute
to shortly acuminate, costae c. 17 on each side, spreading, rather prominent adaxially
in sicco, petiolules to c.9 mm long, densely pilose; pseudogemmula of rather unfurled
leaflets, densely long-hairy. Thyrse to 50 cm long, narrowly paniculiform,
tomentellous; peduncle c. 18 cm long, branches rather distant, to 10 cm long, squarrose;
ultimate branchlets cymulose, many-flowered; pedicels tomentellous. Calyx c.2 mm
tall, cupular, sericeous, margin truncate to weakly 4-lobed. Petals 4 or 5, c. 11 mm
long, c. 1.8 mm wide, linear-spathulate, sericeous without, glabrous within, imbricate,
apices concave (corolla clavate in males). Staminal tube swollen at mouth, weakly
adherent to petals at base, hairy at least along interlobe sutures near apex without,
hairy within, margin with 6—8 linear, more or less 2- or 3-toothed, lobes to 2.5 mm
long; anthers 5—8(9), c. 2 mm long, oblong, locellate, basifixed, slightly exserted.
Disc obscure. Ovary 4-locular, sericeous; style sericeous in proximal 5/6; stylehead
subcylindrical with flattened apex. Infructescences to 30 cm long, axillary or on
twigs behind leaves. Capsule spherical, to at least 9 cm diameter, reddish brown,
latex white; pedicel to 9 mm across. Seeds unknown.
Other material seen: SABAH: Kinabatangan, Tedong, 21 July 1962, Singh SAN 31087
(K!,L!,SAN!,SAR!); Lahad Datu, Kretam, 28 May 1954, Wood SANA 4769 (KEP!,
L!, SING!), Lahad Datu, road to Taliwas, Danum Valley, 24 June 2000, Madani et al.
SAN 143512 (SAN!); Ranau, Lohan F.R., 28 Oct 1963, Pennington 7934 (FHO!, K!,
KEP!, L!, SAN!); Sandakan, Sepilok, boundary between Compts 4 & 5,22 Oct 1963,
Pennington 7915 (FHO!, L!, SAN! = Meijer SAN 39141 (SAN!)), Sg. Daghat 1960
Coupe, 18 Oct 1961, Meijer SAN 27861 (SAN!), “Keratam lineated land’, 27 May
1983, Sigin & Lideh SAN 97058 (SAN!); Tawau, Kalabakan, 30 miles WNW of Tawau,
24 April 1954, Wood SAN A 3696 (KEP!, L!, SING!). KALIMANTAN: Loa Haur,
Figure 1.
Chisocheton maxilla-pisticis. A, flowering twig; B, young leafy shoot, C, older leaflet; D,
adaxial view distal part of staminal tube; E, flower bud; F, longitudinal section of flower bud;
G, abaxial view of distal part of staminal tube; H, fruit. (A from SAN 31087, B from SAN
136830, C from Pennington 7915, D—-G from SAN 31087,H from SAN A 4769.)
191
Chisocheton and Walsura
192 Gard. Bull. Singapore 55 (2003)
west of Samarinda, 60 m, 12 May 1952, Kostermans 6834 (A!, K!, KEP!, L!); Nunukan
(northern part), 2 Dec 1953, Kostermans 8897 (K!, L!, SING!); W Kutei, Kelindjau
River, 12 June 1954, Kostermans 9571 (K!,L!,SING!), bb 16168 (SING!); SE Borneo,
Tidoeng vicinity, 15 m, bb 17894 (L!); Berouw, 400 m, 4 July 1934, bb 19239, (L!).
PHILIPPINES: Palawan, Pagdanan Range, Ibangley Brookside Hill, 21 Apr. 1984,
Ridsdale SMH 496 (KEP!).
Habitat: Lowland dipterocarp forest.
Notes: Some older sterile and fruiting specimens were previously referred with a ‘?’
(Mabberley, 1979: 346) to the closely allied Chisocheton macrophyllus King, which
does not occur in the Tree Flora area. The specific epithet, meaning shark-jaw, refers
to the open pseudogemmula characteristic of the leaf of this tree.
2. Chisocheton velutinus Mabb., sp. nov., Fig. 2
Chisocheton, C. erythrocarpum affinis, sed inflorescentia longiore facile separanda
est. In ser. Sandoricocarpis indumento velutino distincta est. Typus: Wong 1536,
Brunei, Temburong Dist., Bt. Belalong, west ridge from summit, 650 m, 23 July 1989
(holo SAN!; iso KEP!, SAR!, SING!).
Tree to 25 m tall; bole to 35 cm diameter with small buttresses. Bark smooth, medium
brown to greyish, hoop-marked, inner bark pinkish to reddish brown; sapwood straw;
buttresses to 1.3 m x 7.5 cm. Leafy twigs 1.2—2.5 cm diameter. Leaves to 1 m long,
pseudogemmulate; petioles 10-15 cm, more or less angled, like rachis, fulvo-
velutinous; leaflets in up to at least 12 pairs, oblong, to 24 x 9 cm, minutely pubescent
on sunken midrib, densely velutinous on abaxial surface, bases asymmetrical, rounded,
apices shortly acuminate, lateral veins 15—17 on each side, spreading, rather prominent
adaxially in sicco, petiolules to 9 mm long. Thyrse to 40 cm long, paniculiform,
velutinous; branches rather distant, to 13 cm long; ultimate branchlets cymulose,
few-flowered; pedicels densely pubescent. Calyx c. 6 mm tall, cupular, pubescent,
green, margin very obscurely 4-lobed to truncate. Petals 4(5), linear-spathulate, to 16
mm long, imbricate, puberulous to pubescent without, glabrous within, cream.
Staminal tube cream, weakly adherent to petals at base, more or less hairy without,
margin with 6—8 2-toothed lobes to 1 mm long; anthers 6—8, oblong, c. 3 mm long,
Figure 2.
Chisocheton velutinus. A, flowering twig; B, flower buds; C, longitudinal section of flower
bud; D, abaxial view of staminal tube; E, adaxial view of staminal tube; F, young new shoot;
G, older leaflet; H, fruit. (A—E from Wong 1536, F—H from Kirkup et al. 940.) —
193
Chisocheton and Walsura
ss =
a
mene ELC =F
~ -~-+4e4-7
—-TF
194 Gard. Bull. Singapore 55 (2003)
locellate, basifixed, apices within the tube. Disc obscure. Ovary 4-locular, sericeous;
style more or less glabrous; stylehead shortly cylindrical. Infructescences to 30 cm
long, axillary or on twigs behind leaves. Capsule globose, c. 5 cm diameter (immature),
rostrate when young, velutinous, yellow turning red.
Other material seen: Restricted to Borneo. SARAWAK: Ulu Luak, Sg. Setap road,
Miri, 15 May 1964, S 21307 (K!, SAR!, SING!); Kapit, Ulu Merit, Ng. Sebatong, 6
Oct 1969, S 28793 (K!, SAR!, SING!). BRUNEI: Temburong, Bt. Belitun, 29 Jan
1994, Kirkup et al. 940 (K!, KEP!, SAN!, SAR!, SING!). KALIMANTAN:
Balikpapan, Kostermans 10024 (L!); Peak of Balikpapan (G. Beratus), sandstone,
700 m,9 July 1952, Kostermans 7383 (K!, SING!); Sg. Wain region, N of Balikpapan,
30 m, Kostermans 4175 (SING!).
Habitat: Rain forest at altitudes to 650 m.
Notes: Some of the older fruiting specimens cited above (Kostermans 7383,
Kostermans 10024 and S 28793) were tentatively placed under Chisocheton species
B (Mabberley 1979: 372; 1995: 186). The remaining specimens from the Tree Flora
region cited there (S$ 25844 from Kapit, Melinau, Ulu Sapurau, Bt. Salong, Sarawak
(K!,SAR!), with flowers are in bud) is the flowering element used in that description
and remains the only flowering material I have seen of that tree. Apparently conspecific
with it is the fruiting specimen S 4551/5 (from the 7" Division, Belaga, Linau, Sg.
Iban in Sarawak (FHO!, L!, SAR!), collected as C. medusae).
3. Chisocheton sp. nov. aff. diversifolius Mig.
Collected as Aglaia affinis Merr. [= A. odoratissima Blume], this fruiting collection
of a 4 m tall treelet (PC. Yii S 48450, Sarawak, 7" Division, Batang Balleh, Bukit
Melatai, Camp 2; FHO!, SAR!) has delicate twigs with 7-jugate leaves, with very
narrow multi-veined leaflets 5—6 times as long as wide, and a very delicate fruiting
rachis c. 20 cm long with pubescent spherical fruits 3-4 cm diameter. It appears to be
close to C. diversifolius Miq. from Sumatra, but that rarely collected treelet is known
for certain only in flower and has broader leaflets (see Mabberley 1979: 358; 1995:
174).
4. Chisocheton nicobarianus Debnath & Sreek.
The only other species described in the genus since the author’s monograph, now
almost a quarter of a century old, is Chisocheton nicobarianus Debnath & Sreek.
Chisocheton and Walsura 195
from the Nicobars. Examination of an isotype shows it to be conspecific with C.
macrophyllus King:
Chisocheton macrophyllus King in J. Asiat. Soc. Bengal 64(2) (1895) 32; Mabb. in
Bull. Brit. Mus. Nat. Hist. Bot. 6 (1979) 345. Type: (lectotype [selected here from
King’s syntypes] Curtis 2469, Peninsular Malaysia, Penang, ‘Polo Boeting’ (CAL!);
isolectotypes BM!, K!, K [ex SING]!). Synonym: C. nicobarianus Debnath & Sreek.
in J. Econ. Tax. Bot. 16 (1992) 553. Type: Dwivedi 8541, Gt Nicobar, Laful forest, 18
May 1981 (holo CAL, n.v.; iso L!, PBL n.v.). syn. nov.
Note: This brings the total number of species in the genus to 53, with 22 in Borneo
(11 restricted to that island), all of which are found in the Tree Flora area.
Walsura Roxb.
In his revision of the genus Walsura, Clark (1994: 249, 279) discussed the variation
in his concept of W. pinnata Hassk. and proposed informal entities for the major
variants he recognised. In Sabah, these are the typical ‘pinnata’, represented by SAN
86994 (SAN!, SAR!) from Keningau in Ranau district, and also his ‘villamilii’
represented by SAN 57288 (SAN!, SAR!) from Lahad Datu, Sandakan, which has
leaves with seven or nine, as opposed to five, leaflets with some costae failing to
reach the leaf margin, but otherwise intergrading with ‘pinnata’, the form found
throughout most of the range of this tree which, according to Clark (1994: 279, fig.
12) is distributed from the mainland SE Asia to Peninsular Malaysia, Borneo, the
Philippines and NW New Guinea.
Although there is a wide range of costa number in the leaflets of Sabah
specimens, the great majority fall into Clark’s broadly circumscribed W. pinnata,
though specimens with a very small number of leaflet costae (6 or 7 as opposed to
14—19) and short inflorescences and infructescences (e.g. SAN 30551 and SAN 31339;
both in SAN!, SAR!) from Tawau, and SAN 44553 (SAN!, SAR!) from Keningau,
seem rather distinctive, at least within the Tree Flora area.
Moreover there is a small number of other sheets included by Clark and these
are more clearly separable and indeed represent a distinct species described below.
1. Walsura decipiens Mabb., sp. nov., Fig. 3
Walsura, W. trichostemon Migq. affinis sed foliolis ovatis-acuminatis glabris facile
distinguenda est. Typus: Meijer SAN 36201 , Sabah, Labuk Rd mile 25.5 (holo SAN!;
iso K!, KEP!, L!, SAR!, SING! [duplicate (‘Burseraceae’) also sent to BO, n.v.]).
Synonym: W. pinnata Hassk. sensu Clark (1994: 276, 300), p.p. & (1995: 48), p.p.
196 Gard. Bull. Singapore 55 (2003)
Tree to 28 m tall with bole to 6 m, 30 cm diameter. Bark scaly, outer bark dark
reddish brown, inner bark pinkish red. Young twigs pale brown, finely sericeous,
lenticellate, leafy ones c.4—5 mm diameter; apical buds strongly fulvous-hairy. Leaves
to 25 cm long, 2-jugate; petiole 3—5 cm long, terete; leaflets 5.5—14.5 x 2.0-4.5 cm,
the apical larger than laterals, ovate, articulated at petiolule apices, glabrous, somewhat
glaucous abaxially in sicco, base acute, apex acuminate, costae 8-12 on each side,
arising almost at right angles to midrib but strongly arcuate, looping together but not
reaching margin; petiolules 8-18 mm long, that of apical leaflet to 25 mm long,
conspicuously swollen at both ends. Inflorescences subcorymbose cymes to 20 cm
long, in axils of current flush of which apical leaves scarcely developed, all axes
densely fulvous pilose; peduncle to 7 cm long, with branches to 9 cm long, bearing
apical head of branchlets, each branched once or twice more and bearing cymules of
1—5 flowers; bracts to 2 mm long, usually much less, triangular, densely hairy,
caducous; pedicels c. | mm long; bracteoles 0.5—1 mm, triangular, densely hairy,
persistent. Calyx c. 0.8 mm high, green, lobes broadly triangular, apices rounded to
acuminate, very hairy without. Petals oblong, c. 3 mm long, c. 1 mm wide, white,
hairy without, apex obtuse. Stamens 10, with the filaments alternately long and short,
the longer almost as long as petals, weakly bifid, apically strigose, pale green; anthers
c. 0.7 mm long, ovate, apiculate, sparsely hairy, bright yellow, inserted between 2
apical lobes. Disc fleshy, annular. Ovary glabrous; style to 1.2 mm long; stylehead
flattened with apical papilla and peripheral flange. Fruits ellipsoid, 2.5—3 cm long,
brownish green tomentellous, apex usually apiculate. Seed 1, ellipsoid.
Other material seen: Restricted to the northern parts of Borneo. SABAH: Nabawan,
Sg. Millian, 7 Nov 1986, Krispinus SAN 118575 (L!, SAN!); Sandakan, Lungmanis
Timber Camp, 9 Aug 1962, Mikil SAN 31560 (KEP!, L!, SAN!, SAR!); Tawau mile
12.5 Kalabakan road, 25 July 1962 (first record), Aban Gibot SAN 30552 (KEP!, L!,
SAN!, SAR!); Keningau, Saang Lian logging area, LANAS, 28 Oct 1986, Krispinus
SAN 118523 (E!, K!, KEP!, SAN!). SARAWAK: Baram, Batang Tinjar, Ulu Sg.
Sekiwa, Long Kerangan, | Sept 1974, Tong S 35017 (KEP!, L!, SAN!, SAR!).
Habitat: Rain forest, at least sometimes on sandstone, to 150 m.
Notes: The only flowering material known to me is that of the type.
Plants of this species have misled many field collectors who have referred
Figure 3.
Walsura decipiens. A, leafy twig; B, flowering twig; C, flower bud; D, open flower; E, open
flower with some sepals and petals removed; F, stamens; G, fruiting twig; H, cross-section of
fruit; I, indumentum of fruit wall; J, indumentum of lower leaflet surface. (A from SAN 31560,
B-F from SAN 36201, G—J from S 35017.)
197
Chisocheton and Walsura
198 Gard. Bull. Singapore 55 (2003)
them variously to Dialium (Leguminosae), Burseraceae and to several genera in the
Sapindaceae, curators filing them variously (and even in Lauraceae), so the wide
deception is commemorated in the specific epithet.
2. Walsura grandifolia Rid.
In Sarawak, ‘typical’ Walsura pinnata has rarely been collected, though forms with a
wide range of costa number (see above) are preserved (e.g. S 64519 from Kuching
(SAR) and §$ 23959 (SAR) [18 or 19 costae], Pennington 8000 [8 or 9] (SAR) from
Kapit), but S 350/7 from Baram District included by Clark in his broad concept of
the species is here referred to W. decipiens (see above).
Some other Sarawak sheets included by Clark constitute another discrete
entity worthy of specific rank, much as he has afforded the striking stout-shooted W.
pachycaulon Mabb. ex T. Clark. (Clark, 1994). These distinctive specimens, with
very large rugose leaflets and long inflorescences, were collected from treelets
apparently restricted to limestone and they include the type of W. grandifolia Ridl.,
which is here resurrected from Clark’s synonymy, bringing to 16 the number of extant
species of Walsura now recognised, seven in Borneo (to which six are restricted), all
recorded in the Tree Flora area.
Walsura grandifolia difters from W. pinnata not only in its very large leaflets,
inflorescences and flowers, but also in its androecium of almost free filaments, which
have no apical teeth, its larger disc and its longer cylindrical stylehead. In its
androecium therefore it more approaches sect. Surwala (M. Roem.) Hook.f. (see Clark
1994, 1995), such that the distinction between that section and sect. Walsura, to which
W. pinnata belongs, seems hardly worth maintaining. On the other hand, the third
group, sect. Ruswala T. Clark, with the sole species W. dehiscens T. Clark, is marked
by its distinctive dehiscing fruit and, in that, approaches Heynea Roxb.
Walsura grandifolia Ridl. in Bull. Misc. Info. Kew 1930 (1930) 370, g.v. for
description, amplified in Tree Flora account. Type: Kalong in Haviland 1635, Sarawak,
nr Kuching [‘e.s.f.’ = 5 Oct 1892] (holo K!; iso SAR!). Synonym: W. pinnata Hassk.
sensu Clark (1994: 276, 300) & (1995: 48), p.p.
Distribution: Restricted to Kuching Division, Sarawak. Other sheets seen: Teng Bukap,
mile 32 Padawan Road, S 32633 (SAR!); Bt Mentawa, mile 34 Padawan Road, S$
41069 (SAR!); Bt. Bidi, Bau, S 50390 (L!, SAR!); and Bt. Taipu, S 74985 (SAR!).
Habitat: Forest on limestone at altitudes to 220 m.
Notes: Of other specimens referred to Walsura pinnata by Clark (1994: 300), Mogea
Chisocheton and Walsura 199
3711 (L!) is Sandoricum borneense Mig.; SAN 28613 (SAR!) is Aglaia leptantha
Mig. (and, of those referred to Pseudoclausena chrysogyne (Miq.) T. Clark there):
Kokawa & Hotta 3069 (SAN!) = Heynea trijuga Roxb.; S$ 26985 (SAR!) =
Aphanamixis borneensis (Miq.) Harms; S$ 35236 (SAR!) = Micromelum minutum
(Forst.f.) Wight & Arn. [Rutaceae]; and S$ 39029 = Aglaia leptantha [C.M. Pannell,
pers. comm.]). Nonetheless, the rest of the perplexing complex (see diversity of costa-
number and inflorescence size above) around W. pinnata needs further collecting and
analysis in the field across its range, particularly concentrating on whether certain
small flowering trees are really juvenile forms of larger ones, or comprise distinct
species like W. grandifolia. As it stands, even shorn of all the above, W. pinnata still
seems to me to be heterogeneous, and certainly so across its range, a conviction
confirmed by acursory examination of Peninsular Malaysian material now available.
3. Walsura yunnanensis C.Y.Wu
Among his ‘insufficiently known species’, Clark (1994: 290) has W. yunnanensis
C.Y.Wu. Examination of the type preserved at KUN shows it to be W. pinnata sensu
lato (W. cochinchinensis (Baill.) Harms), already known from Yunnan:
Walsura pinnata Hassk., Retzia | (1855) 147, 235, sensu lato; Clark (1994: 276), p.
maj. p. Type (selected by Clark, /.c.): Koorders 971, Java, Bogor, tree no. III b 20
[grown from material coll. Hasskarl, South Bantam, Sept-Oct 1841], 26 June 1892
(neotype BO; ‘clonotype’ Anon. s.n., idem, L [Acc.926258-685]!). Synonym: Walsura
yunnanensis C.Y. Wu, Fl. Yunnanica 1 (1997) 226 cum tab.; Luo ef al. in J. Nat.
Prod. 63 (2002) 947 & Acta Bot. Yunn. 23 (2001) 515. Type: Y.H. Li 2927, Yunnan,
Mengla, highway km. 20-37, Feb. 1961 (holo KUN!), syn. nov.
4. Walsura xizangensis C.Y.Wu & H.Li, referred to Glycosmis Corréa (Rutaceae) by
Clark (1994)on the advice of van Balgooy, has been transferred to that genus as G.
xizangensis (C.Y.Wu & H.Li) D.D.Tao (syn. G. motuoensis D.D.Tao, G. medogensis
D.D.Tao ex P.H.Huang) in Acta Phytotax. Sin. 32 (1994) 369, though Glycosmis is
badly in need of a thorough overhaul.
Acknowledgments
This work was carried out as part of the Tree Flora of Sabah and Sarawak Project (IRPA
Grant 08-04-01-0165/2003). I am greatly indebted to the Director General of FRIM, the Director
of the Sabah Forestry Department, the Director of the Sarawak Forestry Department, and to
Engkik Soepadmo (KEP) for arranging my involvement in the project, and to the staff at the
participating organisations for their kindness and co-operation. I acknowledge the generosity
200 Gard. Bull. Singapore 55 (2003)
of the Keepers and Curators of herbaria (K, KEP, KUN, L, SAN, SAR and SING) for the loan
of specimens and facilities rendered. I am also grateful to Anne Sing (Oxford University
Herbaria) for help in checking a number of points in FHO, Ding Hou (L) with help in reading
Chinese characters, Ruth Kiew (SING) for locating an isotype of Walsura decipiens, and to
Ubaldus Majawal (SAN) and Joseph Pao (SAR) for the illustrations.
References
Clark, T.P. 1994. The species of Walsura and Pseudclausena genus novum (Meliaceae). Blumea.
38: 247-302.
Clark, T.P. 1995. Walsura. Flora Malesiana. 1,12: 45-55.
Mabberley, D.J. 1979. The species of Chisocheton (Meliaceae). Bulletin of the British Museum
(Natural History). Botany Series. 6: 301-386.
Mabberley, D.J. 1995. Chisocheton. Flora Malesiana. I, 12: 136-187.
Gardens’ Bulletin Singapore 55 (2003) 201-208
Arisaema fimbriatum (Araceae) and its
Intraspecific Variation
G. GUSMAN
Université Libre de Bruxelles (CP 232), Faculté des Sciences,
Bd. du Triomphe, B-1050 Bruxelles, Belgium
Abstract
The status of Arisaema fimbriatum Mast. is discussed based on the examination of herbarium
material as well as living plants in the wild and in cultivation. A. fimbriatum is recognised as
having two subspecies: subsp. fimbriatum and subsp. bakerianum (Eng1.) G.Gusman established
here.
Introduction
Arisaema fimbriatum Mast. occurs exclusively on limestone in Peninsular Malaysia
and the southernmost provinces of Thailand, i.e. in the Malesian phytogeographical
region. The species name recalls the spadix-appendix covered with long, thin and
filiform bristles. The crimson and crinate appendage, protruding far out from the
spathe is so outstanding that A. fimbriatum is one of the most remarkable Arisaema
species yet described from this area.
M.T. Masters encountered the plant in autumn 1884 in Sander’s Nursery, St.
Alban’s, England, and described it the same year in The Gardeners’ Chronicle.
In 1911, a new plant was introduced in the trade, first called A. bakeri Hort.
ex Ridley or A. bakerianum (Hort.) Sander, which Engler (1920) eventually described
as A. fimbriatum var. bakerianum in Das Pflanzenreich. Nevertheless, some authors
question the relevance of this variety and treat A. fimbriatum var. bakerianum as
synonymous with A. fimbriatum.
I have gathered new information from examination of herbarium specimens,
living plants in the field in Malaysia and Thailand, and also in cultivation. As a result
of their morphological differences, the two taxa are considered distinct, and, from
their separate geographical distributions, the two varieties of A. fimbriatum are changed
to subspecies.
History
Arisaema fimbriatum was first discovered in cultivation in Sander’s nursery among
orchids collected in the Philippines (Masters, 1884). Later, when reintroduced by
Charles Curtis from Langkawi Islands, its correct Malaysian origin was established
202 Gard. Bull. Singapore 55 (2003)
in the south of Langkawi. Its distribution extends in Peninsular Malaysia from Perlis,
Kedah, Perak, Kelantan, Pahang, as far south as Selangor. In Perlis it is found in the
Tale-Ban National Park, along the Thai border, always growing in crevices in limestone
rocks. Its northernmost limit seems to be in Kuan Nieng, Khao Chang Low, Thailand,
where Kerr collected it in 1928 (Jacobs, 1962) as no specimen of A. fimbriatum has
been found further north of Kerr’s location.
Gagnepain (1941), apparently unaware of Master’s species, described Kerr’s
specimen as A. putii Gagnep. Hetterscheid & G.Gusman (2003) have shown that the
three syntypes of A. putii cited by Gagnepain include at least two different species!
This confusion was recently resolved and the lectotypification of Kerr 15899 (K)
established the synonymy of A. putii with A. fimbriatum.
Arisaema fimbriatum subsp. bakerianum was first collected by Curtis in 1896
during a boat trip along the coast from Penang to Kasom (Curtis, 1897). As noted on
Curtis 3281, it was collected in ““Kasoom on limestone island near the mouth of the
river’, today in the Ao Phang Nga National Park.
1. Arisaema fimbriatum Mast. subsp. fimbriatum, Gard. Chron. 22 (1884) 680.
Type: Gard. Chron. 22 (1884) fig. 119.
Synonym: Arisaema putii Gagnep., Notul. Syst. (Paris) 9 (1941) 127. Lectotype: Kerr
15899, THAILAND, Kuan Nieng, Khao Chang Low, 24 July 1928 (K).
Plant deciduous, up to 60 cm tall and wide. Subterranean stem a subglobose tuber,
3-5 cm wide, 2—3.5 cm high, cream. Roots fleshy, few. Pseudostem the length of the
petioles or longer, 25—30 cm long, 1.8 cm wide at the base, pale magenta or pale
green. Cataphylls up to 5, lanceolate, 2—30 cm long, similar in colour to the pseudostem.
Petiole 20-25 cm long, 1 cm wide at the base, similar in colour to the pseudostem
with faint, pink stripes in the upper part. Leaves usually 2, trifoliolate. Leaflets glossy
dark green above, paler with slightly prominent veins beneath; subsessile or shortly
petiolulate; margins entire; apex acuminate ending in a short arista, 0.5 cm long.
Central leaflet elliptic ovate, 20—24 cm long, c. 15 cm wide; base cuneate or convex.
Lateral leaflets ovate, slightly asymmetrical, nearly the size of the central leaflet;
base convex. Inflorescence above the foliage, emerging when the leaves unfold.
Peduncle 20-30 cm long, 6 mm wide at the base, similar in colour to the petioles.
Spathe-tube cylindrical, slender, 6-7 cm long, 1.5—2 cm wide, dark crimson with
pale green stripes outside and white stripes inside; mouth-margins straight to hardly
recurved. Spathe-limb horizontal, ovate lanceolate, 7-10 cm long, 3.54 cm wide,
dark crimson and white striped, with a green hue outside. Spathe-tip acuminate. Spadix-
appendix long exserted from the spathe at anthesis, 8-15 cm long, slender; exserted
portion a crimson, pendulous, whip-like tail, covered with c. 25 mm long, filiform,
Arisaema fimbriatum 203
Figure 1: Arisaema fimbriatum subsp. fimbriatum
A) Flowering plant; B) Spadix appendage; C) Spathe
204 Gard. Bull. Singapore 55 (2003)
dark crimson, bristly projections; included portion light yellow-green, crimson striped,
4—6 mm across at the base; with scattered green neuters, c. 5 mm long; lower part
sessile. Spadix male or bisexual. Male part c. 3 cm long, 0.5 cm across; stamens
loosely arranged, borne on a green stalk, c. 1 mm long, 2—4-androus, anthers crimson;
thecae dehiscent by an elongated pore. Female part c. 2 cm long and 1.5 cm across;
pistils loosely packed; ovaries ovoid and green, stigma stalked and penicillate.
Infructescence subcylindrical, c. 2.2 cm long and 1.8 cm across, borne on an erect
peduncle when ripe; receptacle cream. Berries densely packed. Seeds 1 mm in diam.;
testa brown. Germination a simple eophyll is produced just after germination.
Flowering period: May to August. Ripening time: November.
Distribution: Peninsular Malaysia (Kedah, Kelantan, Pahang, Perak, Perlis,
Selangor) and SW Thailand (Songkhla).
Habitat: Primary, tropical lowland evergreen rain forest, 100-300 m altitude,
growing in rich humus, limestone crevices.
Additional material: MALAYSIA — KEDAH: Langkawi, Curtis 1890 (SING), Curtis
1893 (SING), Moosa 1896 (SING), Bukit Puteh Corner s.n. 20 November 1941
(SING), Pulau Dayang Bunting FRI 44601 7 May 1996 (KEP), Telok Apau Haniff &
Nur SFN 7490 (SING). KELANTAN: Gua Musang UNESCO Limestone Expedition
427, 6 August 1962 (SING). PAHANG: Bukit Sagu Nur SFN 25160, 14 October
1931 (SING); Kota Glanggi Ridley s.n., 1891 (SING). PERLIS: Pulau Rabana
Henderson SFN 23099 (SING). PERAK: 10 km south of Ipoh Ng FRI 1605, 4 October
1966 (SING). SELANGOR: Batu Takun near Kanching Nur SFN 34376, 3 November
1937 (SING).
Notes: FRI 4460] clearly shows that the subterranean stem is tuberous and not
rhizomatous as has sometimes been erroneously stated in the literature (Govaerts &
Frodin, 2002).
2. Arisaema fimbriatum subsp. bakerianum (Engl.) G.Gusman stat. nov.
Type: Curtis 3281, THAILAND, Kasom, “on limestone island near mouth of the
river’, November 1896 (lectotype SING, selected here).
Synonym: A. bakeri Ridl., Journ. Roy. Asiat. Soc. Bengal 59 (1911) 218.A. bakerianum
Engl. Pflanzenr. 73:4, 23F (1920) 152, pro syn. A. fimbriatum var. bakerianum Eng}.
Pflanzenr. 73:4, 23F (1920) 152.
Subspecies bakerianum differs from subsp. fimbriatum by the following characters:
Shoot usually with 1 leaf, rarely 2. Leaflets pale green beneath with a reddish hue
when unfolding. Inflorescence: spathe-tube white-green, unstriped outside and inside;
mouth margins light green; spathe-limb crimson outside paler inside with a white
Arisaema fimbriatum 205
Figure 2: Arisaema fimbriatum subsp. bakerianum
A) Inflorescence; B) Spadix appendage; C) Habitat
206 Gard. Bull. Singapore 55 (2003)
spot at throat level, surrounded by light green margins; spadix-appendix included
portion pale yellow-green and glabrous.
Distribution: SW Thailand, endemic on islands in Ao Phang Nga (Bay of Phang Nga)
and Ao Phra Nang.
Habitat: North facing slopes in limestone crevices, beneath bushes, just above sea
level.
Additional material: THAILAND. From islands facing Kasom: Curtis 3262,
November 1896 (SING); Curtis 3281, November 1896 (BM, SING); Curtis s.n., 1897,
a specimen which flowered in Penang BG in April 1897 (SING); Curtis 8947, 1897
(SING). Krabi, Ao Phra Nang: Gusman 03076, July 2003 (BR).
Notes: Curtis 3281 has a tuberous subterranean stem. Curtis also noted that “A.
fimbriatum has always two leaves, this has one”’, in full agreement with observations
I made in the field. I had the opportunity to observe hundreds of plants, which were
always on islands and near the seashore. By contrast, limestone outcrops on the
mainland along Ao Phang Nga and Ao Phra Nang did not harbour any specimens of
A. fimbriatum. This may result from the coolness the proximity to the sea provides on
these islands.
Discussion
Reliable morphological characters that separate the two subspecies are summarized
in Table 1 and illustrated in Figures 1| and 2.
Table 1. Morphological differences between Arisaema fimbriatum subsp. fimbriatum
and subsp. bakerianum.
CHARACTERS ssid
usually 2 usually |
Number of leaves
Colour of the crimson with white crimson without
spathe-limb stripes stripes
Colour of the crimson with white white-green without
spathe-tube stripes stripes
Arisaema fimbriatum 207
In addition, there are clear differences in both geographic distribution and
habitat. Their distributions are allopatric, which justifies the modification of status
from variety to subspecies. A. fimbriatum subsp. fimbriatum is widespread on limestone
in Peninsular Malaysia from the Langkawi Islands south to Selangor. Curtis, who
visited the west coast of the Peninsular Malaysia from Penang to Kasom, did not
collect specimens of subsp. fimbriatum north of Langkawi, nor do | know of other
specimens collected north of Songkhla. In contrast, A. fimbriatum subsp. bakerianum
is restricted to islands southwest of the Thai coast, roughly from Kasom to Krabi. A
gap separates the distributions of the subspecies.
Nor do they share the same habitat even though both are always found growing
on limestone rocks. All specimens of subsp. fimbriatum I know were collected on
hills at c. 100 m altitude or above, sometimes quite far from the coast, while I always
found populations of subsp. bakerianum on islands growing on cliffs close to the
shore only a few metres above sandy beaches.
Taxonomic position
Hara (1971) chose Arisaema fimbriatum as the lectotype of Arisaema sect. Fimbriata
Engl. Today, sect. Fimbriata has been revised (G. & L. Gusman, 2002) and includes
only tuberous and deciduous species. The rhizomatous, evergreen species are now
included in the new sect. Anomala G. & L. Gusman (in Hetterscheid & G.Gusman,
2003). All species in sect. Fimbriata are tropical species found in NE India, Myanmar,
Thailand, Cambodia, Vietnam, S China, Malaysia, Indonesia, the Philippines, and
possibly in Laos. All of them have spadix appendages with neuters, at least in female
or bisexual appendages, present at the base and, often also to the part of the appendage
protruding from the spathe-mouth. In most species, these bristles are stiff and fleshy.
In A. fimbriatum, the appendage, outside the spathe-tube, is densely covered with
long, flexuous, bristly filaments. In sect. Fimbriata, the closest species is A. barbatum
Buchet from Java, which has the same kind of hairy appendage. Engler specially
created sect. Barbata for A. barbatum (and A. balansae), a section no longer in use
today. They are easy to separate: namely by their spathes. It is auriculate at the mouth
in A. barbatum; the margins of the spathe-mouth are straight in both subspecies of A.
fimbriatum.
Acknowledgments
I thank David Scherberich, Jardin Botanique de Lyon, France, who provided living
material of A. fimbriatum subsp. fimbriatum from the Langkawi Islands, Phongsawat
Phinhiran and Chanrit Sinhabaedya for material of A. fimbriatum subsp. bakerianum
from S. Thailand. The author is grateful to Leng-Guan Saw (KEP) for images of the
KEP specimens and to Ruth Kiew (SING) for images of the SING specimens, and for
208 Gard. Bull. Singapore 55 (2003)
providing valuable information. Thanks also to Simon Mayo (K), Nguyen Van Du
(HN) and Cees Lut (L) for their help during this study and the reviewers for advice on
nomenclature.
References
Curtis, C. 1897. Annual Report of the Botanic Garden Penang. Botanic Gardens
Singapore. p. 11.
Engler, A. 1920. Fimbriata. In: A. Engler (ed.) Das Pflanzenreich 73 (1V, 23F):
151-153. Berlin, Engelman.
Gagnepain, F. 1941. Aracees Nouvelles Indochinoises. In: Notulae Systematicae
(Paris).:9: 127.
Govaerts, R. and D.G. Frodin. 2002. World Checklist and Bibliography of Araceae.
Royal Botanic Gardens Kew, U.K. p. 192.
Gusman, G. and L. Gusman. 2002. The Genus Arisaema. A.R. Gantner Verlag KG,
Ruggell, Lichtenstein.
Hara, H. 1971. A revision of the eastern Himalayan species of the genus Arisaema
(Araceae). Bulletin of the University Museum, University of Tokyo. 2: 324.
Hetterscheid, W.L.A. and G. Gusman. 2003. Three new Arisaema species from
Thailand and some taxonomic/nomenclatural notes. Aroideana. 26: 33-42
Hooker, J.D. 1890. Arisaema fimbriatum. Native of the Malay Islands. Botanical
Magazine. 116: t. 7150.
Hooker, J.D. 1894. Arisaema. Flora of British India 6: 502.
Jacobs, M. 1962. Reliquiae Kerrianae. Blumea. 9: 462-463.
Masters, M. T. 1884. New Garden Plants. Arisaema fimbriatum, Mast. sp. nov.
Gardeners Chronical n.s.22: 680, 689.
Ridley, H.N. 1911. An account of a botanical expedition to Lower Siam. Aroideae.
Journal of the Royal Asiatic Society of Bengal. 59: 218.
Gardens’ Bulletin Singapore 55 (2003) 209-218
A Revision of Dyera (Apocynaceae: Rauvolfioideae)
DAVID J. MIDDLETON
Harvard University Herbaria
Cambridge, USA
Abstract
The genus Dyera is revised. Two species, Dyera costulata and Dyera polyphylla, are recognised.
A key to the species is presented and descriptions given.
Introduction
This account of Dyera is the latest in a series of revisions of the genera of Apocynaceae
for the forthcoming account of subfamilies Apocynoideae and Rauvolfioideae in Flora
Malesiana. The species of Dyera, under the name jelutong or its spelling variants, are
familiar to many people in western Malesia: Dyera costulata (Miq.) Hook,f. is a
widespread forest tree which is sometimes also cultivated, and Dyera polyphylla (Miq.)
Steenis is a common tree in peat swamp forests. Even though these species are
reasonably well known, the purpose of this revision is to provide a key to and
descriptions of these two species, clarify the nomenclatural issues and typify the names.
Monachino (1946) provided a detailed discussion of the traditional and economic
uses of Dyera and an extensive bibliography of the previous literature. He summarised
the taxonomic information on the genus at that time but stated that his work was to be
regarded as provisional. He did, however, provide a great deal of useful information
on what was known on the species of Dyera up to 1946 including wood anatomy,
commercial yields, silviculture and field observations. I have provided distribution
maps in this work but, as Monachino indicates and I can confirm from my own
observations, most of the trees are only rarely in flower and are, therefore, rarely
collected. The distribution maps must be considered an indication of the range of
distribution of the species rather than the precise localities in which they can be found.
Monachino (1946) and others have suggested that the distinction between
the two species is not clear. Monachino stated: “I emphasize that it is highly dubious
whether D. lowii [= D. polyphylla| merits specific separation from D. costulata.” He
does, however, maintain it, a fate that he does not accord the then often recognised D.
laxiflora, considered by him and by me as a synonyn of D. costulata. Although the
two species recognised here are closely related, in my view their delimitation is not
as difficult as has been hitherto suggested. I have found that the number of specimens
that are not clearly assignable to one species or the other are very few and that the two
210 Gard. Bull. Singapore 55 (2003)
species differ in several quite distinct characters (see key). In addition, Monachino
(1946) suggested that the distinction based on altitude is misleading as D. costulata is
also found at the lower altitudes characteristic of D. polyphylla. However, the issue is
not altitude per se but whether the habitat 1s peat swamp forest or not. This is a habitat
largely confined to lower altitudes but not all low altitude forests are in peat swamps.
D. polyphylla is found only in peat swamp forest and has not been collected above 30
m altitude, whereas D. costulata is found in a wider variety of forests and on different
soils up to 1220 m altitude but at lower altitudes never in peat swamps.
Monachino (1946) provided a long bibliography for the literature on Dyera.
Below I have cited the literature from 1946 or earlier only if it is directly concerned
with the taxonomy of Dyera and the reader is directed to Monachino’s work for further
references.
Herbarium material was studied from the following herbaria: A, BKF, BM,
E, GH, K, KEP, L, NY, P, SAN, SAR, SING, U, US (Holmgren et al., 1990). All
specimens cited have been seen unless otherwise stated. The dimensions given in
the descriptions are for dried material except for the gynoecium and androecium
characters which are for flowers rehydrated with water.
Dyera Hook.
J. Linn. Soc. 19 (1882) 293; K. Schumann in Engler & Prantl, Nat. Pflanzenfam. 4, 2
(1895) 139; Monachino, Lloydia 9 (1946) 182; Pichon, Mém. Mus. Nat. Hist. Nat.
sér. 2,27 (1948) 190.
Trees, often growing to enormous size; buttresses absent; white latex present in all
plant parts. Branchlets usually strongly longitudinally ribbed when young,
occasionally weakly so. Leaves verticillate; petioles relatively long with noticeable
intrapetiolar stipules; blade often weakly crenulate at margin, glabrous. Inflorescence
of umbelliform or paniculate cymes, lax or somewhat congested in upper parts. Sepals
with colleters inside, connate at base, lobes often of different sizes. Corolla lobes
overlapping to the left in bud; mature corolla platter-shaped, lobes oblong to lanceolate,
more or less symmetrical, auriculate at the base on both sides, glabrous outside and
inside. Stamens free from the pistil head, completely included in the corolla tube;
filaments short and narrow; anthers lanceolate, base cordate, apex apiculate, sterile at
apex, dehiscing laterally. Disc annular, inconspicuous, adnate to the ovary. Ovary
apocarpous but carpels closely associated and appearing syncarpous, pubescent; ovules
many per carpel; style and pistil head short. Fruit of paired, divergent follicles, these
heavy and woody, dehiscing at maturity. Seeds elliptic, flattened, with a broadly
membranous wing.
Dyera (Apocynaceae) 211
Key to the species of Dyera
Trees without pneumatophore roots; leaf blades mostly obtuse to shortly acuminate
at apex, only rarely rounded (and then not exclusively so), broadly cuneate to
SOC AURIS ISR! Johor. ol Ses WA RN es. G8 1. D. costulata
Trees with pneumatophore roots; leaf blades emarginate at apex, only rarely with
some (but not all) leaves on a plant with rounded or apiculate apices, cuneate and
pomuibicht mabe peti at. bases ty Yet 202) ARLE 2. D. polyphylla
1. Dyera costulata (Mig.) Hook.f. (Map 1)
Dyera costulata (Miq.) Hook.f., J. Linn. Soc. 19 (1882) 293; Hook,f., Fl. Brit. Ind. 3
(1882) 644; King & Gamble, J. As. Soc. Beng. 74, 2 (1907) 443; Winkler, Bot.
Jahrb. 49 (1913) 372; Merrill, Bibl. Enum. Born. Pl. (1921) 498; Ridley, Fl. Mal.
Pen. 2 (1923) 345; Kerr in Craib, Fl. Siam. Enum. 2 (1939) 441; Masamune,
Enum. Phan. Born. (1942) 619; Monachino, Lloydia 9 (1946) 190; Browne, Forest
Trees of Sarawak and Brunei (1955) 60; Backer & Bakhuizen f, Fl. Java 2 (1965)
227; Smythies, Common Sarawak Trees (1965) 21; Whitmore, Tree Fl. Mal. 2
(1973) 13; Cockburn, Trees Sabah | (1976) 19; Anderson, Check]. Trees Sarawak
(1980) 148; Whitmore & Tantra, Checkl. Sumatra (1986) 19; Ashton, Man. non-
Dipt. Trees Sarawak (1988) 32; Corner, Wayside Trees Mal. ed. 3, 1 (1988) 154;
Whitmore & Tantra, Checkl. Sulawesi (1989) 14; Whitmore, Tantra & Sutisna,
Checkl. Kalimantan. (1990) 25; Kessler & Sidiyasa, Trees Balikpapan-Samarinda.
(1994) 53; Turner, Gard. Bull. Sing. 47 (1997) 12; PROSEA 5, 2 (1995) 229;
Coode et al. (eds.), Checkl. Pl. Brunei (1996) 26; Middleton in Argent et al.,
Man. Non-Dipt. Trees Central Kalimantan | (1997) 83; Kochummen, Tree FI.
Pasoh For. (1997) 151; Middleton, Fl. Thailand 7(1) (1999) 36; PROSEA 18
(2000) 65. Basionym: A/stonia costulata Migq., Fl. Ind. Bat. Suppl. (1861) 556.
Type: Diepenhorst HB 1114 Sumatra, Priaman (lecto U, designated here; iso K,
Lp
Synonyms: Alstonia grandifolia Miq., Fl. Ind. Bat. Suppl. (1861) 555. Type: Teijsmann
HB 4044, Sumatra, Palembang (lecto U, designated here; iso K, L).
Alstonia eximia Miq., Fl. Ind. Bat. Suppl. (1861) 555. Type: Teijsmann HB 3358
Sumatra, Bangka, near Djebus (lecto U, designated here; iso L).
Dyera laxiflora Hook,f., Fl. Brit. Ind. 3 (1882) 644; King & Gamble, J. As. Soc.
Beng. 74, 2 (1907) 444; Ridley, Fl. Mal. Pen. 2 (1923) 345. Type: Cantley 226
Singapore (lecto K, designated here).
Dyera sp. indet. Coode et al. (eds.), Check]. Pl. Brunei (1996) 27.
Tree to 80 m high, to 3 m diameter, sometimes with somewhat exposed roots but not
with knee-shaped pneumatophores. Bark dark grey, brown or black, smooth with
212 Gard. Bull. Singapore 55 (2003)
squarish scales; inner bark cream, pale grey or pale reddish; wood cream or white.
Twigs 3.5—9 mm diameter, glabrous. Leaves in whorls of 4-8; stipules 3-6 mm long;
petiole 2—6.2 cm long, glabrous; blade coriaceous to papery, obovate, oblong or elliptic,
5.542 x 1.8-14 cm, 1.64.3 times as long as wide, glabrous above and beneath,
glaucous beneath or not; base subcordate to rounded (sometimes from a narrowed
base), rarely cuneate, margin crenulate or weakly crenulate, apex short acuminate to
rounded; midrib sunken to slightly raised above; secondary veins 12-24 pairs, at 45—80°
from midrib, clearly distinguishable from tertiary venation, prominent or flat above,
prominent beneath; tertiary venation reticulate or subscalariform, prominent above
and beneath. Inflorescences arranged in whorls, 4-18 cm long, glabrous, many-
flowered; peduncle 2.5—9.2 cm long; pedicels 1.5—6.5 mm long. Sepals ovate or
orbicular, apex rounded to acute, 1—3 x 0.8—2 mm, I—1.5 times as long as wide, ciliate
or not, glabrous. Corolla white, yellowish green or pinkish yellow; tube 1.1—-3 mm
long, 0.3-0.6 times as long as lobes, glabrous inside and outside; lobes 3—9 x 1.2—2.3
mm, |.7-4.4 times as long as wide, glabrous outside and inside. Stamens inserted at
050.6 mm from corolla base which is 02-036 of tube length; anthers 1.1-1 4 x 0.4—0.5 mm,
2.7—3 times as long as wide, exserted 0Q—0.88 mm from corolla throat. Ovary 0.3-0.6
mm long; style O0-0.2 mm long; pistil head 0.5—0.7 mm long. Fruit 18-40 cm long,
2.54 cm diameter. Seed grains c. 2.5 x 1.5 cm,c.5 x 2 cm with wing.
Habitat and Ecology: In a range of evergreen forest types on brown or yellowish soil,
at altitudes to 1200 m.
Distribution: Southern Thailand (6 collections), Sumatra (24 collections), Peninsular
Malaysia (31 collections), Singapore (2 collections), and Borneo (Brunei, 2 collections;
Kalimantan, 26 collections: Sabah, 32 collections; Sarawak, 10 collections).
Note: Specimens of Dyera costulata from Sumatra more often have a cuneate leaf
base than those from other parts of its range, making it more difficult to distinguish
herbarium material of the two species from Sumatra.
Monachino (1946) and a number of later authors following Monachino listed
Dyera costulata as possibly occurring in Sulawesi. However, I have not seen any
material from Sulawesi and doubt it occurs there. Kessler et a/. (2002) do not include
it in their checklist to the woody plants of Sulawesi. It is also absent from Java.
2. Dyera polyphylla (Mig.) Steenis (Map 2)
Dyera polyphylla (Miq.) Steenis, Blumea 14 (1967) 316; Anderson, Trees Peat Swamp
Forests Sarawak (1972) 23: Cockburn, Trees Sabah 1 (1976) 20: Anderson, Check].
Trees Sarawak (1980) 148; Ashton, Man. non-Dipt. Trees Sarawak (1988) 32;
Dyera (Apocynaceae) 213
Map |. Distribution of Dyera costulata (Miq.) Hook.f.
Whitmore, Tantra & Sutisna, Check]. Kalimantan. (1990) 25; PROSEA 5.2 (1995)
230; Middleton in Argent et al., Man. Non-Dipt. Trees Central Kalimantan. |
(1997) 83. Basionym: Alstonia polyphylla Miq., Fl. Ind. Bat. Suppl. (1861) 556.
Type: Jeijsmann HB 3212 Sumatra, Bangka, near Djebus (holo U; iso BO n.v.,
rE):
Synonyms: Dyera lowii Hook f., J. Linn. Soc. 19 (1882) 293; Merrill, Bibl. Enum.
Born. Pl. (1921) 498; Masamune, Enum. Phan. Born. (1942) 619; Monachino,
Lloydia 9 (1946) 194; Browne, For. Trees Sarawak & Brunei (1955) 63; Smythies,
Common Sarawak Trees (1965) 21; Whitmore & Tantra, Checkl. Sumatra (1986)
19; Coode et al. (eds.), Check]. Pl. Brunei (1996) 26. Type: Beccari 3570 Sarawak
(lecto K, designated here; iso FI n.v., P).
Dyera borneensis Baill., Bull. Soc. Linn. Paris | (1888) 751; Merrill, Bibl. Enum.
Born. Pl. (1921) 498, Masamune, Enum. Phan. Born. (1942) 619. Type: Beccari
3570 Sarawak (holo P).
Tree to 60 m tall, to 2 m diameter; knee-shaped pneumatophore roots present, these
chocolate-brown and with paler lenticels. Bark chocolate-brown or greyish brown
with corky paler lenticels and horizontal ridges; inner bark cream; wood cream. Twigs
11-14 mm diameter, glabrous. Leaves in whorls of 6 — 8; stipules 4.5—5 mm long;
petiole 2.1-4.5 cm long; blade coriaceous or subcoriaceous, obovate, 4-24 x 2.3-10.7
214 Gard. Bull. Singapore 55 (2003)
cm, 1.7-2.9 times as long as wide, glabrous above and beneath, glaucous beneath,
base cuneate, margin not crenulate or only weakly so, apex retuse or, more rarely,
rounded, obtuse or apiculate; midrib flat or slightly raised above; secondary veins
17-32 pairs with 3-12 mm spacing, 65—75° from midrib, more or less straight to
curved ascending, slightly prominent or flat above, not prominent beneath; tertiary
venation reticulate to somewhat scalariform, not prominent above or beneath.
Inflorescences arranged in whorls, 8.5—-14 cm long, glabrous; peduncle 4-11 cm
long, sometimes with reduced leaves at apex; pedicels 1.5—3 mm long. Sepals ovate
to orbicular, apex rounded or obtuse, 1—1.4 x 1—1.4 mm, 0.9-1.2 times as long as
wide, not ciliate, glabrous. Corolla tube 1-3 mm long, 0.35—1 times as long as lobes,
glabrous or, slightly pubescent beneath stamens inside, glabrous outside; lobes 1.84
x 1-1.7 mm, 1.4-3 times as long as wide, glabrous outside and inside, not ciliate.
Stamens inserted at 0.5-0.6 mm from corolla base which is 0.4—0.43 of the tube
length; filaments c. 0.3 mm long; anthers 0.8-1.5 x 0.3-0.4 mm, 2.7-3.8 times as
long as wide, exserted 0.1-0.8 mm from corolla throat. Ovary 0.5-0.9 mm long;
style 0-O.1 mm long; pistil head c. 0.5 mm long. Fruit 22-30 cm long, 1.8-4 cm
diameter. Seed grains | .8—2 x 0.8—1.2 mm, 3.7-4.5 x 1.2—1.5 with wing.
Habitat and Ecology: Known only from peat swamp forest, often in association with
Alstonia pneumatophora Backer ex Den Berger. Both species have pneumatophore
roots that can easily be distinguished in the field by the chocolate-brown bark with
corky paler lenticels and horizontal ridges of Dyera polyphylla and the grey bark
with non-corky horizontal lenticels of Alstonia pneumatophora even when the
connection to the parent tree is not obvious.
Distribution: Sumatra (10 collections) and Borneo (Brunei, 5 collections; Kalimantan,
13 collections; Sabah, 6 collections; Sarawak 10 collections).
Notes: Flowering material of Dyera polyphylla is extremely scarce and, therefore, the
description given for flowers above is likely to be expanded once more material can
be collected. The initial findings suggest that there is a complete overlap in flower
dimensions for the two species of Dyera. However, the scarcity of flowers for D.
polyphylla makes it more difficult to be conclusive when comparing the flower
characters in the genus. I was unable to confirm the differences in sepals and anthers
tentatively suggested by Monachino (1946). However, the two species are very easily
separable on vegetative characters.
In the publication of the new combination of Dyera polyphylla by Van Steenis,
the type specimen was given as “HB 2312 in U, isotype L, BO”. However, HB 2312
is a typographical error for HB 3212.1 have not seen the isotype specimen in Bogor.
Dyera lowii was originally described by J.D. Hooker with a number of
syntypes, Low s.n., Beccari 3570 and Lobb s.n., and although it was clearly named in
Dyera (Apocynaceae) 215
Map 2. Distribution of Dyera polyphylla (Miq.) Steenis
honour of Sir Hugh Low and therefore it would seem appropriate to lectotypify his
collection the only specimen I have seen, from Kew, is sterile. The Beccari collection
mentioned without number in the protologue is Beccari 3570. The Kew specimen of
this collection is fertile and given that there are also duplicates would much better
serve as the lectotype. The Paris duplicate of Beccari 3570 is also the holotype of
Dyera borneensis Baill. This name is legitimate as Dyera lowii contained a number
of syntypes and was not lectotypified before the publication of Dyera borneensis.
The holotype of D. borneensis in Paris can therefore simultaneously serve as an
isolectotype of D. lowii.
References
Holmgren, P.K., N.H. Holmgren & L.C. Barnett. 1990. Index Herbariorum. \APT,
New York.
Kessler, P.J.A.,M.M. Bos, S.E.C. Sierra Daza, A. Kop, L.P.M. Willemse, R. Pitopang
& S.R. Gradstein 2002. Checklist of the Woody Plants of Sulawesi,
Indonesia.Blumea. Supplement 14.
Monachino, J. 1946. A revision of Dyera (Apocynaceae). Lloydia. 9: 174-202
216 Gard. Bull. Singapore 55 (2003)
Acknowledgements
I thank the curators and staff of the herbaria that loaned material or accommodated
my visits. The Tree Flora of Sabah and Sarawak Project partially funded my work at
the KEP, SAN and SAR herbaria in 2002.
Specimens examined
Only the specimens with a clear collector and collector number are listed. (1) = Dyera
costulata; (2) = Dyera polyphylla.
Aban & Toshifumi SAN 60108 (1),95171 (1); Abdullah BRUN 16911 (2); Abu 1772
(1), 2258 (1), 3328 (1); Agama 4414 (1), 38790 (1); Ahmad 10816 (2), 94494 (1),
3885 (1); Allen & Kadim 468 (1); Aloysius & Dewol SAN 73963 (1); Ambri & Arifin
W312 (1), W513 (1); Amiruddin 47 (1); Ampuria SAN 32633 (1); Anderson S 9730
(2), S 28738 (1); Anggana bb 37128 (2); Asah anak Unyong BRUN 3152 (1); Awang
47831 (1); Awang Enjah S 58063 (2).
Beccari 3570 (2); Binideh SAN 58586 (1); Boschproefstation bb. 804 (1), bb 7334
(2), bb 10553 (2), bb 15222 (1), bb 35690 (1), bb 36151 (1); Bujang 30496 (2), 30549
(2); Burley et al. 1408 (1).
Cantley 226 (1); Carroll 546 (2); Castillo & Valderrama 10 (1); Castro A 820 (1);
Cheng FRI 27548 (1), FRI 27908 (1); J. Clemens & M.S. Clemens 502 (1), 21502
(1); Clements SAN 138208 (1); Cuadra A 1296 (2).
Daris 53560 (1); De Hulster 6 (2); Diepenhorst HB 1114 (1), Dumas 1541 (1); Edwards
3894 (2); Edwards 36671 (2), BNB 3893 (2); Egon A 0613 (2); Endert 37E (1), 479
(1), 509 (1); Enggoh 7252 (1).
Fischer 754 (1); Flemmich 29323 (1); Foxworthy 4906 (1).
Gadoh KLU. 1355 (1); Garai 2049 (1); Grashoff 56 (2), 677 (2), 784 (1).
Hamid 971 (1), 4917 (1), 5499 (1), SF 24216 (1); Haviland 2170 (2); Haviland & C.
Hose 3495 (2); Heyne 765 (2); Holttum SFN 9904 (1).
Ibrahim 14382 (1).
Jawa S 65627 (1).
Dyera (Apocynaceae) Zin
Kadir A 3509 (1), KEP 55431 (1); Kawasan SAN 79623 (1); Keith A 7137 (1); Kerr
13909 (1); Kochummen KEP 76696 (1); Kostermans 107A (1), 4171 (1), 10164 (1),
bb 34207 (1), bb 35355 (1); Kunstler 4689 (1).
L.T.S. SAN 64858 (1); Lai et al. S 68565 (1); Lakshnakara 353 (1); Lasan SAN
102531 (1); Leeuwenberg & Rudjiman 13075 (1); Lobb s.n. (2); Low s.n. (2).
Madani SAN 36768 (1); Maingayi 1097 (1); Mat Yatim 26173 (1); Meijer & Wood
SAN 130247 (2); Van Meurs s.n. (2); Mikil SAN 28098 (1), SAN 31802 (1); Mohamad
17169 (1); Mujin & Tuyok SAN 78716 (1).
Native Collector 804 (2); Neth. Ind. For. Service bb 10589 (2); bb 6315 (1), bb 9847
(2), bb 10644 (2), bb 12936 (1), bb 15221 (1), bb 16255 (1), bb 16265 (1), bb 16280
(1), bb 16400 (1), bb 16407 (1), bb 16742 (1), bb 16962 (1), bb 17952 (1), bb 18112
(2), bb 18246 (1), bb 18400 (2), bb 19973 (2), bb 21178 (1), bb 21212 (1), bb 21263
(2), bb 23480 (1), bb 23929 (2), bb 24663 (1), bb 27601 (1), bb 27744 (1), bb 28079
(2), bb 28122 (1), bb 28446 (1), bb 28564 (2), bb 29133 (2), bb 29145 (2), bb 29444
(2), bb 29993 (1), bb 30122 (1), bb 31601 (1), bb 31754 (1), bb 31997 (1), bb 32283
(2); Ngah 21954 (1); Nicholson 22297 (2); van Niel 3966 (2); Nooteboom 5033 (1);
Nur SEN 35475 (1).
Omar 8855 (1); Onggib 7137 (1); Orolfo 4786 (1); Othman Ismawi et al. S 56417 (1).
Paie S 37571 (1); Pickles S 3565 (1); Pilis Malahim SAN 95257 (1).
Raji 55439 (1); Ramos 1912 (1); Richards 1308 (1); Ridley 62 (1), 4922 (1), 5656
(1); Ridsdale PBU195 (1).
Sadau SAN 49557 (1); Saikeh SAN 72224 (1); Sharin 35177 (1); Sinclair 6335 (1);
Sitam 603 (2); Smitinand & Williams 17003 (1), 17004 (1), 17213 (1), 17214 (1);
Soewanda bb. 32655 (2), bb. 36716 (1); Symington C.F.22122 (1).
Tahja 7 (2); Talib Bidin SAN 84691 (1), SAN 84751 (1), SAN 84752 (1); Talip SAN
55688 (1); Tan SAN A 0617 (2); Tantra 75 (2), bb 35858 (2); Tarmiji SAN 73884 (1);
Tarmiji & M. Alexius SAN 79883 (1); Tarmiji & A. Tasan SAN 82940 (1); Teijsmann
HB 3212 (2), HB 3358 (1), HB 4044 (1).
Watson 39633 (2); Winkler 2435 (1); Wong WKM 252 (1); Wyatt-Smith KEP 79326
(2).
Yakim 1988 (1).
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Gardens’ Bulletin Singapore 55 (2003) 219-228
Curcuma codonantha (Zingiberaceae) — A New Species from
the Andaman Islands, India
J. SKORNICKOVA, M. SABU AND M.G. PRASANTHKUMAR
Department of Botany, University of Calicut,
673635 Kerala, India
Abstract
Curcuma codonantha Skornickova, M. Sabu & Prasanthkumar sp. nov. from the Andaman
Islands, India, is described and illustrated.
Introduction
Species delimitation in the genus Curcuma L., Zingiberaceae, is rather complicated
due to superficial similarity of several species, lack of type material, the short duration
of flowering and the necessity of studying fresh material. Approximately 100 species
are distributed in tropical and subtropical Asia with a few species extending to Australia
and the Pacific Islands. However, the economically important or ornamentally
interesting species are found naturalized all over the tropics and elsewhere, because
they are popular among gardeners. Several new species of Curcuma have been
described from Asia during the past five years (Mood & Larsen, 2001; Sirirugsa &
Newman, 2000). To date, in India about 30 species are reported and accepted
(Karthikeyan et al., 1989; Jain & Prakash, 1995; Skornickova & Sabu, 2002). Many
of them were early described by Roxburgh (1810, 1820), while others were described
or reported quite recently (Bhat, 1987; Mangaly & Sabu, 1988; Mangaly & Sabu,
1993; Sabu & Mangaly, 1988; Sivarajan & Balachandran, 1983; Skornickova et al.
2003; Velayudhan et al., 1990 & 1991).
As part of the project “Revision of Indian Zingiberaceae’, we studied Curcuma
extensively in the field in the Andaman Islands resulting in the documentation of ten
species. Previously, only four species had been reported from the Andamans
(Balakrishnan and Bhargava, 1984; Dagar and Singh, 1997; Srivastava 1998). While
exploring the northern part of the Andaman Islands, flowering specimens of a Curcuma
with a lateral spike, oblong-lanceolate, glabrous leaves and prominent bell-shaped
flowers, which were strongly exserted from the bracts, were collected. Critical
examination of fresh material showed that it did not match completely any other
220 Gard. Bull. Singapore 55 (2003)
Indian species of Curcuma. Due to presence of anther spurs the specimen belongs to
subgenus Curcuma (Eucurcuma K. Schum.). After scrutinising original descriptions
of all the Asian species so far known under the subgenus and, after consulting others
working on the genus in Thailand and Myanmar, we are confident that this species is
new. It is described and illustrated below.
Curcuma codonantha Skornickova, M. Sabu & Prasanthkumar, sp. nov.
Curcumae aeruginosae Roxb. similis rhizomatis figura et colore cremeo, foliis
viridibus macula purpurea secus costam carentibus, coma rosiore bracteis apicaliter
macula brunneopurpurea, floribus campanulatis e bracteis fertilibus exsertis differt.
Typus: India, Andaman Islands, North Andaman, Diglipur District, Kalighat, Alt. 15
m, 13° 09' N 92° 57'E, 22.V.2002, Skornickova & Prasanth Kumar 73319 (holo MH;
iso K, CALI, SING).
Figure 1, Plate 1.
Rhizomatous herb, up to 1.50 m tall. Rhizome 3.5—6 x 3-6 cm, sessile tubers present
5—13 cm long, 1.5—2 cm in diam., sometimes almost on the soil surface, light brown
outside, skin glabrous, creamy-yellowish inside, aromatic (camphoraceous), bitter in
taste, scales triangular, papery, brown, glabrous, present on main rhizome and sessile
tubers, in soil usually quickly decaying and leaving scars; root tubers 3—5 x 1.5—2 cm,
elliptic, white inside, on 2—3 mm-thick roots c. 5-20 cm away from main rhizome.
Leafy shoot up to 1.50 m tall, leaves 3-6; pseudostem 20-45 cm long, sheathed by 2—
4 reddish-green bracts (clearly visible only at the beginning of the season, later drying
and decaying), innermost as long as the pseudostem, outer ones gradually smaller in
length, 3-4 cm broad; ligule 3 mm, hyaline, greenish translucent, 2-lobed, hairy along
the margin, hairs 0.5 mm long; petiole 10-30 cm long, winged and gradually tapering
into lamina; lamina lanceolate or oblong-lanceolate, 35—75 x 8—14 cm, green, paler
on the lower surface, margin translucent white, hyaline, 0.5 mm wide; upper surface
mostly glabrous, but slightly hairy at the apical part of the leaf, especially on veins
and near margins, hairs 0.3 mm long, lower surface glabrous; tip acuminate c. 1—1.5
cm hairy, base oblique, attenuate, gradually tapering into 1 .5—2 mm-wide wings along
the petiole. Inflorescence vernal, lateral. Peduncle 14-23 cm, 0.7—1.5 cm diam.,
covered with 4—6 reddish-green sheaths, glabrous, the innermost longest, structurally
similar to the fertile bracts above, outer sheaths gradually smaller, broad 3—4 cm.
Figure 1. Curcuma codonantha. a. Habit; b. Flower (side view); c. Flower (front view); d. Bracteole; e.
Calyx; f. Dorsal corolla lobe; g. Lateral corolla lobe; h. Labellum; i. Lateral staminode; j. Anther (front);
k. Anther (side); 1. Stigma; m. Epigynous glands and ovary; n. Ovary (cross section). Based on the type
material Skornickova & Prasanthkumar 73319. Del. J. Skornickova. |
Curcuma codonantha from the Andaman Islands 221
REN os ,
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222 Gard. Bull. Singapore 55 (2003)
Spike 12—20 x 6-8 cm, formed by c. 30—35 bracts including coma. Coma forming 1/3—1/4
the length of the inflorescence, coma bracts 7-9, c. 5—6 x 2.5—3 cm, bright pink with
darker brownish-violet patch on the tips, lower side almost glabrous, upper side shortly
hairy, hairs 0.2 mm long, lower coma bracts fertile, 3-4 uppermost sterile. Fertile
bracts 3.5 x 44.5 cm, green, tips sometimes tinged with red, upper side very shortly
hairy, lower side quite glabrous. Cincinnus with 4—6 flowers. Bracteoles 1.7—2.5 x
0.6—1.4 cm, hyaline, translucent white, glabrous. Flowers 6 cm long, yellow, longer
than the bracts, exserted |.5—2 cm from the bracts. Calyx 8 mm long, translucent
white, 3-dentate, unilaterally split for about 3 mm, hairy, hairs 0.4 mm long. Corolla
tube 3.2—3.5 cm, towards base light yellow, towards lobes pink, glabrous. Corolla
lobes pink, glabrous, dorsal lobe 2 x 1.3 cm, mucronate, mucro 4-5 mm, lighter in
colour (whitish pink), hairy, hairs 0.3 mm long, lateral lobes 1.5 x 0.9 cm, pink,
glabrous. Labellum 2 x 1.8 cm, emarginate, yellow, deep yellow in the centre with
whitish lines on the outside border (golden median band), middle lobe emarginate, 3
mm deep. Lateral staminodes 12 x 9 mm, light yellow, with short glandular hairs on
inner side. Anther spurred, 8 mm, anther thecae whitish, 4-5 mm long, filament light
yellow, 4 mm long, constricted, 5 mm at base, 2 mm at upper part. Anther spurs 3 mm
long, whitish yellow, divergent, but slightly incurved. Ovary trilocular, 34 x 3 mm,
densely hairy, glabrous at the base, hairs 0.3-0.4 mm long, ovules many. Stigma
exserted c. 1—1.5 mm from between anther thecae, white, ciliate, 1.4 x 1.4 mm.
Epigynous glands 2, yellowish-green, c. 5 mm long, 0.5 mm diam. Fruits not seen.
The plant description is based on observation of material from the type locality
as well as measurements of fully-grown plants in the Calicut University Botanical
Garden. The leafy shoot at the beginning of season, when the plant is flowering, is
about 60—70 cm with only 2—3 leaves. Later in the season, the plant grows to 1.50 m
with about six leaves. The green-reddish sheaths covering the pseudostem are most
prominent at the beginning of the season, later they dry and decay.
Flowering: May to June, inflorescences occur together with the first leaves.
Distribution: So far known only from the type locality. We have also seen this species
in Havelock Island (south part of Andaman Islands), but unfortunately we were not
able to collect it. Since most of the Curcuma species in the Andamans are recent
introductions connected with migration of people from various parts of India to
theAndaman and Nicobar Islands, it is doubtful if this species is native there. Thus, in
future, the species may be found elsewhere in India.
Plate 1. Curcuma codonantha. |. Habit; 2. Detail of inflorescence; 3. Coma (seen from above); 4.
Rhizome with roots and tubers; 5. Detail of flower in subtending bract (side view). All photographs are
of the type material Skornickova & Prasanthkumar 73319. Photo J. Skornickova.
Curcuma codonantha from the Andaman Islands
224
Gard. Bull. Singapore 55 (2003)
Table 1. Comparison of morphological characters of Curcuma codonantha and C.
aeruginosa. (The diagnostic characters are in bold).
Main rhizome c. 3.5—6 x 3-6 cm,
sessile tubers present 5—13 cm,
1.5—2 cm in diameter. Inwardly
creamy-yellowish colour.
Rhizome
Root tubers 3-5 x 1.5—2 cm,
white inside.
| cae C. codonantha sp. nov. C. aeruginosa Roxb.
Main rhizome c. 10 x 5 cm, obovate-
conical, sessile tubers 5-15 cm,
1.5—2.5 in diameter. Inwardly
aerugineous (bluish-green) colour.
Root tubers 2.55.5 x 1.5—2:5,
whitish inside.
Leafy shoot | Leafy shoot up to 1.5 m, 3-6 leaves.
Pseudostem and peduncle sheathed
by reddish-green bracts.
Ligule 3 mm, 2-lobed, hairy along
the margin, hairs 0.5 mm long.
Lamina Lamina up to 35-75 x 8-14 cm,
base oblique, attenuate.
Adaxially plain green with no
coloration, abaxially pale green.
Inflorescence Vernal, lateral, flowering in
May. Peduncle 14—23 cm,
sheathed by reddish-green bracts,
spike c. 12—20 x 6-8 cm. Coma
c. 1/3-1/4 of the inflorescence.
Coma bracts bright pink with
darker brownish-violet patch at
the tips, c. 5—6 x 2.5—3 cm,
lower side almost glabrous,
upper side shortly hairy.
Fertile bracts 3.5 x 44.5 cm,
green, tips sometimes tinged with
red, upper side very shortly hairy,
lower side quite glabrous.
| Cincinnus 4—6 flowers.
Leafy shoot up to 1.9 m, 4-6 leaves.
Pseudostem and peduncle
sheathed by green bracts.
Ligule 2 mm, obscurely 2-lobed,
glabrous.
Lamina up to 40-90 x 10-20 cm,
base attenuate. Adaxially green
with dark purple cloud along
the midrib on the distal half of
the leaf, abaxially pale green.
Vernal, lateral, flowering in April
-May. Peduncle 10-25 cm,
sheathed by green bracts, spike
c. 14-20 x 6-9 cm. Coma c.
1/3 of spike. Coma bracts pink
or light pink, with no obvious
patch at the tips, c. 5—7 x 1.22 cm,
sparsely hairy on both
surfaces, tip mucronate 0.2—-0.3
mm.
Fertile bracts 4.5—-5 x 3.5-4.5 cm,
green with tips tinged with red-
purple, quite glabrous both sides.
Cincinnus 4—6 flowers.
Flowers Flower 6 cm, bell-shaped, well
exserted from the fertile bracts.
Bracteoles |.7—2.5 x 0.6—1.4 cm,
hyaline, translucent white, glabrous.
Flower 5-5.5 cm, same length as
fertile bracts, not exserted.
Bracteoles 1.7—2.5 x 0.7—1.5 cm,
hyaline, whitish translucent, hairy at
the tip.
225
Curcuma codonantha from the Andaman Islands
Table 1. Continued:-
Calyx 8 mm long, unilaterally split
3 mm, translucent white, 3-dentate,
hairy.
Corolla tube 3.2—3.5 cm, glabrous,
towards base light yellow, towards
lobes pink.
Labellum 2 x 1.8 cm, emarginate,
yellow, deep yellow in the centre,
yellow towards margin.
Lateral staminodes 12 x 9 mm, light
yellow.
Anther thecae 4—5 mm, anther spurs
3 mm, whitish-yellow, divergent.
Ovary 3-4 x 3 mm, densely hairy.
Calyx 11-12 mm, dentate,
unilaterally split 4-5 mm, whitish
translucent with pink at the teeth tips
sparsely hairy.
Corolla tube 2.8—3.2 cm, glabrous,
whitish yellow base, pink tinged
towards lobes, lobes of rich pink
colour.
Labellum 1.8—-1.9 x 1.9-2 cm,
emarginate, deep yellow in centre,
lighter towards margin.
Lateral staminodes 14 x 7 mm,
yellow.
Anther thecae 4.5 x 0.7 mm, anther
spurs 3 mm, white, divergent.
Ovary 3.5 x 3 mm, hairy.
Epigynous glands c.5 mm long,
0.5 mm diameter, yellowish-green.
Epigynous glands 4.5—5.5 mm long,
0.7 in diameter, yellowish green.
Habitat: Open moist places along stream banks.
Uses: Bengali settlers at the type locality use this species for the extraction of starch
from the rhizomes. Extraction of starch, commonly known as East Indian arrowroot,
has been reported for many Curcuma species all over India and it seems that people
use any Curcuma species that grows locally.
Vernacular name: Bengalis living near the type locality call this plant Shodhi. The
name Sat’hi or Sotee was mentioned by Roxburgh (1810) and Shuthee (Roxburgh,
1820) as the vernacular name for Curcuma zerumbet Roxb., which is nowadays usually
treated as synonym of C. zedoaria (Christm.) Roscoe. Even though Roxburgh’s
description of C. zerumbet consists of only a few lines, it is obvious that it is not C.
codonantha, since Roxburgh’s C. zerumbet has a purple cloud down the middle of
leaf and the flowers are shorter than their bracts. Also, from our fieldwork in Bengal
over the past few years we have observed that the name Sotee or Shuthee is used for
several different species of Curcuma.
Etymology: The epithet “codonantha” is derived from the Greek words “codon” (bell)
and “anthos” (flower) referring to the peculiar bell shape of the flowers exserted from
the fertile bracts, the most obvious character of this species.
226 Gard. Bull. Singapore 55 (2003)
Notes: The closest species to Curcuma codonantha is C. aeruginosa Roxb., which is
commonly distributed throughout Kerala and is found in several localitites in the
Andaman Islands. Distinguishing these species in the field might be difficult due to
similar habit and coloration of flower parts, both have a deep yellow labellum and
lateral staminodes with pink corolla lobes, and lateral spikes and they flower at the
same time. The most obvious difference between them lies in the flowers, which in C.
codonantha are bell-shaped and well exserted from fertile bracts, while they are the
same size as the fertile bracts and do not extend beyond them in C. aeruginosa. They
can also be told apart by the rhizome, which in C. codonantha is inwardly creamy-
yellowish in contrast to the inwardly aerugineous (bluish-green colour) of C.
aeruginosa; C. codonantha leaves are plain green, quite glabrous excepting on the
prominent veins and margin areas at the distal half on the upper side of the lamina,
which are sparsely hairy, while C. aeruginosa leaves are glabrous, have a deep purple
cloud, which runs along the both sides of midrib and which is especially prominent at
the distal half of lamina; and the coma bracts of C. codonantha are marked at the tips
by a brownish-violet or red brownish patch unlike of those of C. aeruginosa, which
are pink or light pink and although they can be somewhat darker at the tips, but do not
have a prominent patch of such a deep colour (Table 1).
According to our observations in the field, Curcuma species that do not set
seed and reproduce exclusively vegetatively by sessile tubers are uniform within a
population as well as between populations. Characters like the red patch on the leaves,
indumentum and coloration of flower parts or the inner colour of the rhizome, which
may vary in the case of seed-setting species, are stable for non seed-setting species.
Acknowledgements
The authors thank the Department of Science and Technology, Govt. of India for
financial support (Order No. SP/SO/A-20/99 dt. 09.11.2001). We are also indebted to
staff of Forest Department of the Andaman Islands for their help and the authorities
of Ministry of Environment and Forests for granting permission for collecting. The
senior author thanks the Indian Council for Cultural Relations New Delhi, India and
the Ministry of Education of the Czech Republic for awarding a research fellowship;
to Hlavkova Nadace, Czech Republic for partial financial support for travelling; and
to Singapore Botanic Gardens for providing facilities during her repeated visits. We
thank Dr. J. F. Veldkamp, National Herbarium of the Netherlands, Leiden, for latinising
the diagnosis and to Dr. Axel D. Poulsen for comments on the manuscript.
Curcuma codonantha from the Andaman Islands 227
References
Balakrishnan, N.P. and N. Bhargava. 1984. The genus Curcuma L. (Zingiberaceae)
on Andaman and Nicobar Islands. Journal of Bombay Natural History Society.
81: 510-514.
Bhat, K.G. 1987. Curcuma oligantha Trimen (Zingiberaceae) — a new record for
India. Indian Journal of Forestry. 10: 66-68.
Dagar, J.C. and N.T. Singh. 1997. Plant resources of the Andaman and Nicobar Islands.
(Enumeratio and Illustrations of Vascular Plants). Vol. 11. Bishen Singh Mahendra
Pal Singh, Dehra Dun, India. Pp. 874-875.
Jain, S.K. and V. Prakash. 1995. Zingiberaceae in India: Phytogeography and
endemism. Rheedea. 5: 154-169.
Karthikeyan, S., S.K. Jain, M.P. Nayar and M. Sanjappa. 1989. Florae Indicae.
Enumeratio Monocotyledoneae. Botanical Survey of India, Calcutta. Pp. 289-
PEN
Mangaly, J.K. and M. Sabu. 1988. A new species of Curcuma from South India.
Notes from the Royal Botanic Garden Edinburgh. 45: 429-431.
Mangaly, J.K. and M. Sabu. 1993. A taxonomic revision of the South Indian species
of Curcuma L. (Zingiberaceae). Rheedea. 3: 139-171.
Mood, J. and K. Larsen. 2001. New Curcumas from South East Asia. The New
Plantsman. 8: 207-217.
Roxburgh, W. 1810. Descriptions of several of the Monandrous Plants of India. Asiatic
Researches. 11: 318-362.
Roxburgh, W. 1820. Monandria Monogynia. Flora Indica. Mission Press.
Serampore, India. Pp. 1-84.
Sabu, M. and J.K. Mangaly. 1988. Curcuma vamana (Zingiberaceae): A new species
from South India. Journal of Economic and Taxonomic Botany. 12: 307-309.
Sirirugsa, P. and M. Newman. 2000. A new species of Curcuma L. (Zingiberaceae)
from S.E. Asia. The New Plantsman. 6: 196-197.
Sivarajan, V.V. and I. Balachandran. 1983. A new species of Curcuma from Southern
India. Notes from the Royal Botanic Garden Edinburgh. 41: 321-323.
228 Gard. Bull. Singapore 55 (2003)
Skornickova, J. and M. Sabu. 2002. The genus Curcuma L. in India: Resume and
future prospects. In: A.P. Das (Ed.) Perspectives of Biology, Bishen Singh,
Mahendra Pal Singh, Dehradun, India. Pp. 45-51.
Skornickova, J.,M.Sabu. and M.G. Prasanthkumar. 2003.A new species of Curcuma
L. (Zingiberaceae) from Mizoram, India. Gardens Bulletin Singapore. 55: 89-95.
Srivastava, S.K. 1998. Zingiberaceae in Andaman and Nicobar Islands, India. Higher
Plants of the Indian Subcontinent. 8: 1-33.
Velayudhan K.C., V.S. Pillai and V.A. Amalaraj. 1990. Curcuma kudagensis
(Zingiberaceae) - A new species from Karnataka. Journal of Economic and
Taxonomic Botany. 14: 476-479.
Velayudhan K.C., V.A. Amalraj and V.K. Muralidharan. 1991. Curcuma
thalakaveriensis sp. nov. (Zingiberaceae) - a new species from Karnataka state,
India. Journal of Economic and Taxonomic Botany. 15: 479-481.
Gardens’ Bulletin Singapore 55 (2003) 229-238
Three new species of Porterandia
(Rubiaceae) from Mount Kinabalu, Borneo
M.S. ZAHID
Institute of Biological Sciences,
University of Malaya,
50603 Kuala Lumpur, Malaysia
Abstract
Three new species of Porterandia (Rubiaceae), P. beamanii, P. chanii and P. puffii, which
occur on Mount Kinabalu, Borneo, are described.
Introduction
In the earlier enumerations of plants on Mount Kinabalu, Stapf (1894) and Gibbs
(1914) did not list any species that could be identified as Porterandia, a genus described
by Ridley in 1939. As material accumulated, the Bornean taxa were wrongly named
Porterandia anisophylla from Peninsular Malaysia and Sumatra in herbarium
identification. A revision of the genus has been carried out at the University of Malaya,
where a number of new species have been recognized for Borneo. These include
three undescribed species from Mount Kinabalu. The new species are described here
in anticipation of the forthcoming Volume 5 of The Plants of Mount Kinabalu by
Professor John Beaman’s team. A full revision of Porterandia, consisting of 22 species
in all, will be published subsequently.
The new species
1. Porterandia beamanii MS Zahid, sp. nov. Fig. 1
Porterandiae minori Ridl. similis sed foliis typice minoribius (ad 5-6 cm latis) lobisque
calycis florum femineorum foliaceis in statu fructifero differt. Typus: Pereira et al.
JTP 144 Sabah, Tambunan, road to Trusmadi, lower montane forest, 1180 m (9 Mar
1995, fruiting) (holotypus SAN!, isotypus SING).
Gynodioecious or possibly gynomonoecious trees, to 35 m high, to 25 cm diameter,
not buttressed. Bark smooth to slightly fissured to grid-cracked; pale brown to dark
brown. Shoot tips, distal branch internodes, petioles and leaf veins with erect-suberect
230 Gard. Bull. Singapore 55 (2003)
hairs. Stipules ovate-triangular and fused along the edges to form a tube, 1—1.5 cm
long, densely hairy. Mature leaves with petiole (0.4—)0.5—1.5(—2) cm long, 1-3 mm
thick; lamina usually elliptic, rarely obovate, 8—20.5 cm long, 3—6(—8.5) cm wide;
leaf base cuneate; leaf apex acute to acuminate to short-caudate; when dry chartaceous;
secondary veins 9—15 pairs, on upper side flattened to slightly raised, on lower side
distinctly prominent; tertiary venation a much-branched network between pairs of
secondary veins.
Bisexual inflorescences: Peduncle 0.2—0.6 cm long; habit compact, with 1—2 distinct
branching orders. Flowers 5—21 per cyme, usually in 1—3 clusters; pedicels 1-3 mm
long, 1.5—2 mm thick; calyx tube/limb densely covered by hairs (most of calyx surface
hidden); calyx lobes short to narrow-triangular to linear, often also spathulate becoming
foliaceous in fruit, 2.5—7 mm long; corolla hypocrateriform, the tube 5—8 mm long,
2—3 mm wide at the throat, with a slightly to conspicuous inflated part just below the
throat, outer surface totally covered by upward-pointing hairs; corolla lobes ovate to
narrowly elliptic, 4-5 mm long; anthers 4-5 mm long, with pollen; style 4-5 mm
long, stigma 1—2 mm long. Female inflorescences: Peduncle 0.2—0.7 cm long; habit
sparsely branched, with | distinct branching order. Flowers 1—3 per cyme; pedicels
2—3 mm long, 1.5—2 mm thick; calyx tube/limb densely covered by hairs (most of
calyx surface hidden); calyx lobes narrowly triangular to linear to spathulate becoming
foliaceous in fruit, 3-8 mm long; corolla hypocrateriform, the tube 5—9 mm long, 2—
3 mm wide at the throat, with a slightly inflated part just below the throat, outer
surface totally covered by upward-pointing hairs; corolla lobes narrowly elliptic to
ovate, 4-5 mm long; anthers 4—5 mm long, without pollen; style 4-5 mm long, stigma
3—4 mm long. Fruit sub-globose to ellipsoid, 2—3.5 cm long, 2—3 cm wide, short-
tomentose when young, becoming glabrous. Seeds flattened, lens-shaped to rounded,
4—6 mm x 5—7 mm, testa surface finely areolate.
Notes: Bisexual and female cymes can occur together on the same branch (Ampuria
SAN 32690).
Distribution: Borneo, so far known only from Sabah (including Mt Kinabalu and the
Crocker Range) and Sarawak, in lower montane forest at 3700—6000 ft [1147-1860
m] elevation.
Specimens Examined: BORNEO. SABAH: Ranau, Kundasang, 4300 ft [1333 m],
Singh SAN 28254, 10 Nov 1961, female inflorescences (BO! L!), bisexual inflorescences
Figure 1. Porterandia beamanii. A, leafy branch with bisexual inflorescences. B, bisexual flower,
longitudinal section. C, leafy female fruiting branch. Note foliaceous calyx lobes in B and C.
A from Pereira JTP 141 (SAN), B from Singh SAN 28254 (SAR) and C from Pereira JTP 144 (SAN).
ol
Porterandia from Mount Kinabalu
S
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AO
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232 Gard. Bull. Singapore 55 (2003)
(KEP! SAN! SAR!), Kinabalu National Park headquarters, 5300 ft [1643 m], Aban
SAN 57718, 13 Mar 1967, fruiting (KEP! SAN!), Mount Kinabalu, Ulu Liwagu and
Ulu Mesilau, Chew, Corner & Stainton RSNB 2661 ,2 Apr 1961, female inflorescences
(K! L! ), trail from Kiau Nulu to Marai Parai, near Tohubang River, 3700 ft[1147 m],
Wong WKM 2366, 12 Sep 1993, fruiting (KEP! SAN! SING), Penibukan, 4000-5000
ft [1240—1550 m], Clemens & Clemens 30492 ,28 Dec 1932, bisexual inflorescences
(K! L! SING), Gurulau Spur, 6000 ft [1860 m], Clemens & Clemens 50573, 30 Nov
1933, female inflorescences (K! L!); Ranau, Tenompok FR, Wong WKM 2876, 30
Aug 2001, female inflorescences (KLU! SAN!); Tambunan, Crocker Range, Km 62.5
on Kota Kinabalu-Tambunan road, Beaman 7183,9 Oct 1983, fruiting (L!), road to
Trusmadi, 1180 m, Pereira et al. JTP 141, bisexual inflorescences (SAN! SING) &
JTP 144,9 Mar 1995, fruiting (holotype SAN! isotype SING), Mount Trus Madi,
1200 m, Nooteboom 1451, 20 Mar 1969, fruiting (L! SAN!); Penampang, Crocker
Range, Km 51.8 on Kota Kinabalu-Tambunan road, Beaman 8945, 16 Mar 1984,
fruiting (L!); Keningau, Ulu Sg Tinagalan FR, Fidilis & Asik SAN 113143,22 Nov
1985, bisexual inflorescences (KEP! SAN!); Sandakan, Mamahat Camp, K.F.L, 4000
ft [1240 m], Ampuria SAN 32690, 1 Dec 1962, female inflorescences (L!), bisexual
inflorescences (KEP!). SARAWAK: 4th Division, proposed Gunung Murud National
Park, along Sungai Taramusu, 1500 m, Yii S 44635, 14 Sep 1982, fruiting (K! KEP!
L!), Kapit, Melinau, Ulu Sampurau, Bukit Sampadai, 1433 m, /lias § 40721, 6 Apr
1980, fruiting (KEP! SAN! SAR!).
This species is named after Professor John Beaman, who is well known for his work
on the enumeration of Mount Kinabalu plants.
2. Porterandia chanii MS Zahid, sp. nov. Fig. 2
Porterandiae anisophyllae (Jack ex Roxb.) Ridl. similis sed limbo calycis sparsim
piloso et tubo corollae tecto pilis sursum directis differt. Typus: Singh SAN 34727
Sabah, Sandakan, along north boundary Sepilok FR (1 May 1962, bisexual
inflorescences) (holotypus SAN! isotypi BO! L! SAR! SING).
Gynodioecious tree, to 16 m high, to 18 cm diameter, not buttressed. Bark smooth
with transverse lenticels to slight grid-cracked; pale brown to dark grey. Shoot tips,
distal branch internodes, petioles and leaf veins with erect-suberect hairs. Stipules
ovate-triangular and fused along the edges to form a tube, 1.5—2 cm long, sparse to
densely hairy. Mature leaves (not immature leaves near the shoot-tips) with petiole
Figure 2. Porterandia chanii. A, leafy branch with bisexual inflorescences. B, mature bisexual flower,
longitudinal section. Note very sparsely hairy calyx.
A from Singh SAN 34727 (L), B from Agama A 525 (K).
233
Porterandia from Mount Kinabalu
234 Gard. Bull. Singapore 55 (2003)
1—2.5(—3) cm long, 1-4 mm thick; lamina mostly obovate to elliptic, 16-42 cm long,
8—19 cm wide; leaf base cuneate; leaf apex acute to obtuse to caudate; when dry
chartaceous; secondary veins 16—21 pairs, on upper side flattened to slightly raised,
on lower side distinctly prominent; tertiary venation a much-branched network between
pairs of secondary veins.
Bisexual inflorescences: Peduncle 0.3—0.6 cm long; habit laxly branched, rarely
compact, with 4—5(—8) distinct branching orders. Flowers 12—34(—45) per cyme,
usually in 1—3 clusters; pedicels 3-6 mm long, about 1 mm thick; calyx tube/limb
sparsely covered by hairs (much of calyx surface visible); calyx lobes triangular to
linear, 0.5—2 mm long; corolla hypocrateriform, the tube 11-15 mm long, 3-4 mm
wide at the throat, without any conspicuous inflated part just below the throat, outer
surface totally covered by upward-pointing hairs; corolla lobes narrowly elliptic to
ovate, 7—10 mm long; anthers 5—6 mm long, with pollen; style 8—10 mm long, stigma
3—4 mm long. Female inflorescences: Peduncle 0.3—0.7 cm long; habit sparsely
branched, with 2(—3) distinct branching orders. Flowers 5—12 per cyme; pedicels 3—
7 mm long, 1—2 mm thick; calyx tube/limb sparsely covered by hairs (much of calyx
surface visible); calyx lobes triangular to linear, 0.5—2 mm long; corolla
hypocrateriform, the tube 10-12 mm long, 2-4 mm wide at the throat, without any
conspicuous inflated part just below the throat, outer surface totally covered by upward-
pointing hairs; corolla lobes narrowly elliptic to ovate, 7—9 mm long; anthers 3-4
mm long, without pollen; style 7-8 mm long, stigma 3-4 mm long. Fruit sub-globose
to ellipsoid, 3-4 cm long, 3—3.5 cm wide, sparsely hairy when young, becoming
glabrous. Seeds flattened, lens-shaped to rounded, 3—5 mm oe 3—7 mm, testa surface
finely areolate.
Distribution: North and northeast Borneo, in the Sabah-Tarakan region, in primary
and disturbed lowland forest, c. 100—1800 ft [30-560 m].
Specimens Examined: BORNEO. SABAH: Kudat, Bandau, Tagabu F.R., 1800 ft [558
m], Mait & Anthony SAN 37667, 16 Oct 1963, fruiting (SAN!); Kota Belud, Bukit
Matindok, 1000 ft [310 m], Lajangah SAN 32158, 3 Oct 1962, female inflorescences
(SAR!), bisexual inflorescences (BO! KEP! SAN!); Kota Marudu, Marak-Parak, Ag.
Amin & Mancus SAN 118990, | Apr 1987, fruiting (KEP! SAN!); Ranau, west of Kg.
Nabutan, Aban SAN 94542, 22 Mar 1982, fruiting (KEP! SAN!), Kinabalu, Kg.
Melangkap Tamis, Sg. Kepungit, Lorence LL 669, 4 Jul 1995, fruiting (KEP!
SAN!);Tenom, Crocker Range FR, Aban SAN 66787, 18 Apr 1970, bisexual
inflorescences (K! L! SAR! SING); Nabawan, Sg. Millian, Sumbing SAN 118626, 12
Nov 1986, bisexual inflorescences (SAN!); Sipitang, near Mt. Muruk Miau, Pius &
Soinin SAN 143167, 13 Apr 2000, fruiting (SAN!); Sandakan, Kabili F.R.,; compt. 14,
Porterandia from Mount Kinabalu 2B5
Agama A 525, 21 Oct 1947, bisexual inflorescences (BKF! K! KEP!), along north
boundary of Sepilok F.R., Singh SAN 34727, | May 1962, bisexual inflorescences
(holotype SAN! isotypes BO! KEP! L! SAR!); Kinabatangan, Bukit Mansuli, 200 ft
[62 m], John & Markus SAN 144631, 16 May 2002, bisexual inflorescences (KLU!
SAN!); Lahad Datu, Danum Valley, plot 1, Campbell SAN 112095, 12 Jun 1986,
fruiting (SAN!); Tawau, east of Sg. Serudong, 100 ft [31 m], Bakar SAN 26154, 6
Sep 1961, fruiting (SAN!); Kalabakan, Gunung Tembuku, Fedilis & Sumbing SAN
91472, 8 Mar 1980, fruiting (KEP! SAN!). KALIMANTAN: Nunukan Island, 30 m,
Kostermans 9182, 31 Dec 1953, bisexual inflorescences (K! L!).
This species commemorates Datuk C.L. Chan, who has provided field support for my
study of this genus and much encouragement in my learning of Bornean botany.
3. Porterandia puffii MS Zahid, sp. nov. Fig. 3
Porterandiae scortechinii (King & Gamble) Ridl. similis sed limbo calycis sparsim
tecto pilis brevibus, lobis calycis linearibus 1/3 to 1/2 logitudine limbi calycis, et
tubo corollae tecto pilis sursum directis differt. Typus: Clemens & Clemens 33834
Sabah, Mount Kinabalu, Colombon River (Jun 1933, bisexual inflorescences)
(holotypus L! isotypus K!).
Gynodioecious tree, to 15 m high, to 16 cm diameter, not buttressed. Bark grid-cracked;
greyish white. Shoot tips, distal branch internodes, petioles and leaf veins with sparse
appressed hairs. Stipules ovate-triangular and fused along the edges to form a tube,
0.6—1.5 cm long, sparsely hairy. Mature leaves (not immature leaves near the shoot-
tips) with petiole 1—2.3 cm long, 1—1.5 mm thick; lamina mostly obovate to elliptic,
10.5—15(—19) cm long, 4.5—7(—10.5) cm wide; leaf base cuneate; leaf apex broadly
acute to obtuse with a short point; when dry chartaceous; secondary veins 10—15
pairs, on upper side flattened to slightly raised, on lower side distinctly prominent;
tertiary venation a much-branched network between pairs of secondary veins.
Bisexual inflorescences: Peduncle 0.2—0.3 cm long; habit compact, with 1—2 distinct
branching orders. Flowers (2—)6—10 per cyme, usually in 1—2 clusters; pedicels 3—6
mm long, 1—1.5 mm thick; calyx tube/limb sparsely covered by hairs (much of calyx
surface visible); calyx lobes narrow and linear, 2—5 mm long; corolla hypocrateriform,
the tube 8—12 mm long, 4—5 mm wide at the throat, with a conspicuous inflated part
just below the throat, outer surface totally covered by upward-pointing hairs; corolla
lobes ovate-elliptic, 6-9 mm long; anthers 5—6 mm long, with pollen; style 4-6 mm
long, stigma 4—5 mm long. Female inflorescences (in young flower-bud stage):
Peduncle 0.1—0.3 cm long; habit unbranched. Flowers | per cyme; pedicel 3—5 mm
long, 1—1.5 mm thick; calyx tube/limb sparsely covered by hairs (much of calyx
surface visible); calyx lobes narrowly triangular, 2—4 mm long; corolla hypocrateriform,
236 Gard. Bull. Singapore 55 (2003)
the tube with a conspicuous inflated part just below the throat, outer surface totally
covered by upward-pointing hairs; corolla lobes ovate-elliptic; anthers not known.
Fruit sub-globose to ellipsoid, 2.7—-4.8 cm long, 2.6—4.5 cm wide, sparsely hairy
when young, becoming glabrous. Seeds flattened, lens-shaped to rounded, 5—6 mm x
6—8 mm, testa surface finely areolate.
Distribution: Sabah (on Mount Kinabalu and the Crocker Range) and Sarawak, in
lower montane to mossy forest, c. 5|000—6560 ft [1550—2000 m].
Specimens Examined: BORNEO. SABAH: Ranau, mile 35 Ranau road, 5000 ft [1550
m], Aban SAN 57767, 10 May 1967, bisexual inflorescences (KEP! SAN!), Mount
Kinabalu, Mesilau River, Chew & Corner RSNB 4873, 8 Apr 1964, fruiting (K! L!),
Colombon River, Clemens & Clemens 33834, Jun 1933, bisexual inflorescences
(holotype L! isotype K!), 1 mile north of Tenompok on path to Lumu Lumu, Wood A
4463,18 Jul 1954, bisexual inflorescences (L!); Tambunan, road to G. Alab, Pereira
etal. JTP 118,3 Mar 1995, female inflorescences (SAN!). SARAWAK: 4th Division,
Kelabit Highland, summit of Apad Runan, 2000 m, Yii S 56231, 10 May 1988, bisexual
inflorescences (SAN!).
This species is named for Professor Christian Puff, whose work on the Rubiaceae is
well known and who has given much help and encouragement to my own studies.
Acknowledgements
I thank the Keepers and Curators of the herbaria at the Herbarium Bogoriense (BO),
Kew (K), Forest Research Institute Malaysia at Kepong (KEP), University of Malaya
(KLU), Leiden (L), Forest Research Centre, Sandakan (SAN), Forest Research Centre,
Sarawak (SAR) and the Royal Forest Department, Thailand (BKF) for loans of material
for study. Dr. Y.F. Lee and Ms. Joan Pereira of the Forest Research Centre, Sandakan
and Datuk C.L. Chan kindly provided logistic support during my fieldwork. Dr. R.C.K.
Chung of the Forest Research Institute Malaysia assisted while at KEP. Professor
A.L. Lim and Dr. K.M. Wong (University of Malaya) provided much guidance, advice
and encouragement in the preparation of this paper. Professor Christian Puff of the
Institute of Botany, University of Vienna provided helpful suggestions and the Latin
diagnoses. This paper results from an M.Sc. programme at the Faculty of Science,
University of Malaya, and was supported in part by the Malaysian IRPA Project No.
09-02-03-0090-EA090.
Figure 3. Porterandia puffii. A, leafy branch with bisexual inflorescences. B, bisexual flower, longitudinal
section. Note sparse appressed hairs on petioles and leaf veins.
A from Clemens & Clemens 33834 (L), B from Wood A 4463 (L).
Porterandia from Mount Kinabalu 237
2cm
238 Gard. Bull. Singapore 55 (2003)
References
Gibbs, L.S. 1914. A contribution to the flora and plant formations of Mount Kinabalu
and the highlands of British North Borneo. Journal of the Linnean Society,
London, Botany. 42(285): 1-240, pls. 1-8.
Ridley, H.N. 1939. Notes on some Malayan Rubiaceae. Gardenieae. Bulletin of
Miscellaneous Information, Royal Botanic Gardens, Kew. 593-597.
Stapf, O. 1894. On the flora of Mount Kinabalu in North Borneo. Transactions of the
Linnean Society, London, 2nd Series, Botany. 4: 69-263, pls. 11-20.
Gardens’ Bulletin Singapore 55 (2003) 239-256
The Significance of Pollen Morphology in the Taxonomy of
Grewia and Microcos (Tiliaceae) in Peninsular Malaysia and
Borneo
R.C.K. CHUNG', E. SOEPADMO! AND A. L. LIM?
'Forest Research Institute Malaysia
Kepong, 52109 Kuala Lumpur, Malaysia
“Institute of Biological Sciences, University of Malaya
50603 Kuala Lumpur, Malaysia
Abstract
The pollen morphology of 5 species of Grewia and 31 species of Microcos (including their
type species) from Peninsular Malaysia and Borneo were studied using light, scanning and
transmission electron microscopy. The pollen of Grewia species differs from that of Microcos
in being larger with a mean polar axis of 53—75 um and an average equatorial diameter of 39—
55 um, having a coarsely reticulate tectum, a lumina diameter of more than 1.5 um and a
mean exine thickness of 1.55—2.15 um. Microcos pollen is characterised by its smaller size
with a mean polar axis of 25—35 um and an average equatorial diameter of 19-27 um, a finely
reticulate tectum, a lumina diameter of less than 1.5 um and a mean exine thickness 0.84—0.90
um. The pollen of all species examined conforms to the previously recognised ‘Grewia-type’.
Introduction
The genus Grewia consists of about 280-300 species of trees, shrubs or climbers,
distributed from Madagascar, tropical Africa northwards and southeastwards to the
Himalayas, China and Taiwan, India, Sri Lanka, Myanmar, Thailand, Indo-China,
Malesia, Western Pacific and the northern parts of Australia. In the Malesian region
about 30 species are known, of which four occur in Peninsular Malaysia and Borneo.
Microcos is a genus of about 80 species of trees and shrubs, occurring in tropical
Africa (not Madagascar) and Indo-Malesia. In Malesia, some 42 species are known,
of which 31 occur in Peninsular Malaysia and Borneo (Chung, 2001; Bayer and
Kubitzki, 2003).
Previous studies on members of the order Malvales showed that pollen
morphological characters were useful for segregating the Tiliaceae from the
Bombacaceae, Malvaceae and Sterculiaceae (Erdtman, 1952; Nair, 1962; Chaudhuri,
1965; Chaudhuri and Mallik, 1965; Sharma, 1969; Coetzee and Van Der Schijff, 1978;
Martinez-Hernandez et al., 1978; Zhang and Chen, 1984; Sudhakar and Rao, 1987).
240 Gard. Bull. Singapore 55 (2003)
Palynological literature on the family Tiliaceae has been reviewed by Erdtman
(1952). He described over 70 species of the family, including some Peninsular
Malaysian and Bornean representatives, distributed over 45 genera. Since then there
have been a number studies on the pollen of Grewia and/or Microcos from India
(Sharma, 1969; Sudhakar and Rao, 1987), China (Long, 1982; Zhang and Chen, 1984;
Wang et al., 1995) and Sudan (El Ghazali, 1993). These studies show the Tiliaceae to
be a eurypalynous family. Based mainly on aperture type and using tectum
ornamentation as subsidiary characters, Sharma (1969) recognised 7 major and 18
minor pollen types, while Zhang and Chen (1984) recognised four pollen types and
Erdtman (1952) noted only three pollen types in the Tiliaceae.
Erdtman (1952) classified the pollen of Grewia as belonging to a single pollen
type, i.e. the ‘“Grewia-type’ characterised by tricolporate grains with reticulate tectum.
He, however, did not examine the pollen of Microcos. Later, Long (1982), Zhang and
Chen (1984) and Wang et al. (1995) categorised pollen of Grewia and Microcos as
belonging to the ‘Grewia-type’. However, Sharma (1969), using characters of the
tectum ornamentation and lumina width, distinguished four pollen types in the Indian
species of Grewia.
Based on pollen morphological data, Erdtman (1952), Zhang and Chen (1984),
Christensen (1986), and Nilsson and Robyns (1986) concluded that the pollen
morphology of Tiliaceae closely resembled that of Bombacaceae and Sterculiaceae
but differed from that of Malvaceae. Zhang and Chen (1984) supported the separation
of Microcos from Grewia in the Tiliaceae. Sharma (1969) suggested that pollen
morphological characters, when combined with other morphological data, can provide
useful additional taxonomic characters for delimitating genera and species of the
Tiliaceae.
The aim of this study is to investigate the morphological variation in the pollen
of Grewia and Microcos; to assess the taxonomic value of the pollen in the Peninsular
Malaysian and Bornean species; and to contribute to a better understanding of the
pollen morphology of both genera.
Materials and methods
Pollen used in the present study was obtained from herbarium specimens listed in
Table 1.
The pollen samples were acetolysed following the method of Erdtman (1960).
The acetolysed pollen samples were divided for light microscopy (LM) and scanning
electron microscopy (SEM) studies. For LM observations, pollen grains were mounted
in glycerine jelly stained with safranin and sealed with paraffin wax. Measurements
of polar length, equatorial width, colpus length and width were carried out using an
Olympus BX41 microscope. For each sample, 20 pollen grains were measured using
a calibrated eyepiece micrometer. For SEM observations, the acetolysed pollen grains
Pollen morphology of Grewia and Microcos
Table 1. Source of materials studied.
Species®
Grewia
G. occidentalis L. (type species)
G. huluperakensis 1.M. Turner
G. laevigata Vahl
G. multiflora Juss.
G. polygama Roxb.
Microcos
M. paniculata L. (type species)
M. antidesmifolia (King) Burret
var. antidesmifolia
M. antidesmifolia (King) Burret
var. hirsuta (King) Burret
M. borneensis Burret
M. cinnamomifolia Burret
M. dulitensis Airy Shaw
M. erythrocarpa (Ridl.) Airy Shaw
M. fibrocarpa (Mast.) Burret
M. globulifera (Mast.) Burret
M. gracilis Stapf ex Ridl.
M. henrici (Baker f.) Burret
ssp. acuta R.C.K.Chung
M. hirsuta (Korth.) Burret
M. kinabaluensis R.C.K.Chung
M. lanceolata (Miq.) Burret
M. latifolia Burret
241
Collector
Herbarium
Moss 3959"; Relinhoonk 4837
Turner 94-94"
Hashim SAN 33470*; Kostermans
5234"; Lee S. 41948*; Sigin et al.
SAN 110601*; Stone 11856*;
Whitmore FRI 3729*
Motley 240°
Kiah SFN 35286’; Symington
FMS 467167
Cramer 3405’; Kanodia 88480°
Cockburn FRI 7816’; King’s
Collector 4029*; Whitmore FRI
S417"
Creagh s.n. *; Elmer 20911*; Sibat
ee
Haviland 2837*; Tiggi S. 3311
Anderson S. 25147*; Charington
SAN 29852’; Coode MC 7716?
Richards 1767’
Kochummen FRI 16417*; Mohd.
Shah MS 2757
Kochummen KEP 80640’:
Maingay 1080*; Mohd. Shah and
Sidek MS 1164’
Maingay 3064*; Ogata KEP
105036*
Chai SAN 26066; Coode et al.
MC 6776; Haviland 1683
Ahmad and Termiji SAN 67973°;
Ilias S. 45121*; Kostermans
10424?
Clemens 27284*; Haviland 1471*;
Korthals s.n. (L sheet no.
944 56.122) #
Chew and Corner RSNB 4994?:
Sadau SAN 49708?
Chelliah KEP 104621°; Mahmud
FMS 3717*
King’s Collector 7762*; Sarawak
Museum Series 165*; Wong
WKEM 1545*
BM; L
SING
SAN; SING;
KEP; SAN;
KLU; KEP
K
KEP KEP
L; BM
KEP; K; KEP
K; SING;
KEP
SAR; SAR
SAR; KEP;
KEP
SING
KEP; SING
REer: ih:
SING
Kk KEP
SAN; KEP;
SAR
SAN; KEP; L
SING; SING;
IE
SAN; SAN
KEP; SING
K; SAR;
SING
242
Table 1 continued
Species®
M. latistipulata (Ridl.) Burret
var. latistipulata
M. laurifolia (Hook. ex Mast.) Burrett
M. malayana R.C.K.Chung
membranifolia R.C.K.Chung
opaca (Korth.) Burret
ossea Burret
pachyphylla Merr.
pearsonii (Merr.) Burret
phaneroneura Burret
reticulata Rid.
stylocarpoides Burret
subcordifolia R.C.K.Chung
subepetala Stapf ex Ridl.
sumatrana (Baker f.) Burret
tomentosa Sm.
M. triflora (Blanco) R.C.K.Chung
var. triflora
M. triflora (Blanco) R.C.K.Chung
var. longipetiolata (Merr.)
R.C.K.Chung
“used for LM and SEM
“used for LM, SEM and TEM
riparia (Boerl. & Koord.) Burret
Gard. Bull. Singapore 55 (2003)
Collector
Burkill SFN 7826’; Elmer 20310*;
Teo TP 391?
Kochummen FRI 16338*;
Zainuddin FRI 17887*
Loh FRI 19249’; Symington and
Kiah SFN 28797?
Singh SAN 30672*; Yii S. 61492’
Korthals s.n. (L sheet no.
908.253.752)*; Lee S. 432277
Clemens 21071*; Haviland 42’;
Wong WKM 1290?
Aban SAN 35880*; Salleh et al.
BRUN 15315’
Singh and Aban SAN 30135’;
Wright S. 29113?
Hallier 2868"; Ilias S. 281287
Ashton S. 17800*; Creagh s.n. *
Hallier 2208"; Yeob FMS 3190?
Burley et al. NGS 3243*; Othman
S. 37444#
Kirkup and Thomas DK 727’;
Niga NN 118%
Haviland 1885*
Fedilis SAN 95781*; Forbes 2684*
Ang FRI 21599*; Suppiah FRI
28239": Zainuddin and Latiff AZ
|Pa He
Anon. PNH 35319*; Tai SAN
58042?
Singh SAN 30375*; Suib S.
23458*
Herbarium
SING; L;
SING
KEP; KEP
KEP; SING
KEP; KEP
L; SAR
SAR; SING;
BRUN
SAN; KEP
SAN; SAR
BO; SAR
KEP; K
BO; KEP
KEP; SAR
KEP; KEP
K
KEP- 1.
KEP; KEP;
KEP
KEP; SING
KEP; KEP
* Microcos crassifolia Burret without any flowering material was excluded from this study
Pollen morphology of Grewia and Microcos 243
were dehydrated through an ethanol series and critical point dried by using liquid
CO, as the exchange medium. After coating with gold, they were observed with a Jeol
JSM-6400 scanning electron microscope. For transmission electron microscopy (TEM)
studies, the pollen samples were fixed in 1% OsO, for 90 min, dehydrated through an
ethanol series, and then embedded in epoxy resin. Ultrathin sections, cut with a glass
knive, were stained with 8% uranyl acetate for 5 minutes and then Reynold’s lead
citrate (Reynolds, 1963) for 10 minutes. Observations were made using a Philips
CM12 TEM.
Pollen morphological data for all species are summarised in Table 2. The size
classes of pollen following Erdtman (1952) are classified as small (10—25 um), medium
(25-50 um) or large (50-100 um). Terminology follows that of Erdtman (1952) and
Punt et al. (1994).
Results
GREWIA L.
(Table 2; Plates 1A—C, 2A—C, 3A—D and 4A-B)
Light Microscopy
The pollen grains of Grewia species are single, isopolar, radially symmetric,
tricolporate, and rounded-triangular in equatorial outline (ambs). The pollen grains is
medium to large (44-80 um polar length) with an average polar length of 52.9-74.8
uum and an average equatorial axis 38.6—55 um (Table 2). Pollen grain shape varies
from subprolate (P/E = 1.26) to prolate (P/E = 1.36—1.45).
The apertures are tricolporate. The colpi are 1/2—3/4 of polar axis length, narrow
(c. 1.5 um) to wide (3.5-4.5 um), provided with acute apices and delimited by a
clearly defined margin. The endoapertures are lolongate or lalongate, either as wide
as, or slightly wider than the colpi.
Scanning Electron Microscopy
The pollen grains of Grewia have a bireticulate-tectate exine. The muri of the primary
reticulum are smooth, but somewhat irregular in thickness. The lumina of the primary
reticulum are coarsely reticulate and variable in width and shape, and are infilled by
a secondary reticulum comprising few to many finer reticulations and/or perforations
of varying width and shape. Two types of tectum ornamentation were observed in
Grewia pollen: type C (coarsely reticulate with a few perforations/microreticulations)
and type D (coarsely reticulate with numerous perforations/microreticulations).
Transmission Electron Microscopy
The exine is 1—2.54 um thick and consists of an outer ectexine and an inner endexine.
Gard. Bull. Singapore 55 (2003)
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245
Pollen morphology of Grewia and Microcos
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246 Gard. Bull. Singapore 55 (2003)
The ectexine is thicker than the endexine except at the aperture region. The tectum is
0.23-0.86 um thick. The columellae are simple, straight, distinctly arranged in single
rows in a reticulate pattern, and either support the raised muri of the primary reticulum
or support the sunken secondary reticulum. The columellae are 0.46—1.46 um tall and
0.29-0.77 um thick. The spaces between columellae range 0.47—1.1 Um in height
and 0.18—1 um in width. The foot layer is 0.24—0.53 um thick at the mesocolpia, and
is often thickened near the aperture.
The endexine appears smooth, thin and is not always discernible except at the
aperture region. Below the endexine, at the apertural region, there are generally darkly
stained, stout, radial elements towards the intine. The intine is fibrillar and, particularly
at the apertures, stratified into a fibrillar layer and enveloping a lens-shaped body
with a microtubule layer.
Remarks
Apart from minor differences, the pollen morphological characters of the Peninsular
Malaysian and Bornean species of Grewia conform to those of G. occidentalis (the
type species of the genus), the pollen of G. occidentalis compared with Grewia species
from Peninsular Malaysia and Borneo has a thinner tectum and columellae (0.73-
1.23 um), and foot layer and endexine (0.06—0.2 Um).
The pollen of Grewia laevigata can be distinguished from pollen of other Grewia
species by the perforations of the secondary reticulum that encroach into the margins
of the primary muri. Pollen of G. polygama differs from that of G. huluperakensis
and G. multiflora by having fewer microreticulations/perforations of the lumina.
Sharma (1969) reported that pollen size of Grewia species from India was (27.2-)
31(—78.2) um in polar length and (18.7—)21(—57.8) um in equatorial diameter.
However, among the 17 species he studied, one was a Microcos species: G. microcos
L.1is now Microcos paniculata. By omitting the pollen size of this species (P=(272-)
31(—34) um; E = (18.7—)21(—22.1) um) from the data, the pollen size of these Grewia
species in India was (40.8—)47.4(—78.2) um in polar length and (28 .9—)32.6(—57.8)
uum in equatorial diameter. Therefore, in India pollen size of Grewia species was
larger than that of Microcos species (= G. microcos).
The exine thickness of Grewia species from China was reported by Zhang and
Chen (1984) as more than 2.3 um. This falls within the range of the present study,
0.73-2.54 um thick.
MICROCOS L.
(Tables 2 and 3; Plates 1D—F, 2D—F, 3E—H and 4C—D)
Light Microscopy
The pollen grains in Microcos are single, isopolar, radially symmetric, tricolporate,
and rounded-triangular in equatorial outline (ambs). Size ranges from small to medium
Pollen morphology of Grewia and Microcos 247
(20-42 um polar length) with an average polar length of 25.1—35 um and an average
equatorial diameter of 18.6—26.9 um. Most Microcos species possess medium-sized
pollen grains (Table 2). Pollen grain shape varies from prolate spheroidal (P/E =
1.13) to subprolate (P/E = 1.19—1.32) or prolate (P/E = 1.35-1.62).
The colpi are 1/2 to 3/4 the length of the polar axis, narrow to wide, with acute
apices, and delimited by a distinct margin. In most species the endoapertures are
circular, but are lalongate in M. globulifera and circular to lalongate in M. pachyphylla,
M. pearsonii, M. tomentosa, and M. triflora (both varieties). In most species, the
endoapertures are as wide as or slightly wider than the colpi.
Scanning Electron Microscopy
The tectum of Microcos species are bireticulate-tectate. The muri of the primary
reticulum are smooth and of somewhat irregular thickness. The lumina of the primary
reticulum are finely reticulate and are infilled by secondary reticulum with minute to
large microreticulations/perforations of variable number and shape. Two types of
tectum ornamentation are found in Microcos pollen: type A (finely reticulate with a
few perforations/microreticulations) and type B (finely reticulate with numerous
perforations/microreticulations).
Transmission Electron Microscopy
The exine is 0.70—1.05 um thick and consists of an outer ectexine and an inner
endexine. The ectexine is thicker than nexine except at the aperture region. The tectum
is 0.18—0.46 um thick. The columellae are simple, straight, and distinctly arranged in
single rows in a reticulate pattern, either supporting the raised muri of the primary
reticulum or supporting the sunken secondary reticulum. The columellae are 0.2—0.5
um tall and 0.14—0.32 um thick. The spaces between columellae is 0.18—0.45 um in
height and 0.14—0.55 um in width. The foot layer varies in thickness but is generally
markedly thickened at the aperture margin.
The endexine is usually thin and always traceable except towards the apertures,
where it may be thicker. The intine often has numerous microtubule layers enveloped
by the fibrillar layer. At the apertural region there are relatively stout, darkly stained,
radial elements between intine and endexine.
Remarks
The pollen morphology in the species examined matches that of M. paniculata (the
type species of the genus). The palynological data in this study conform well with the
results reported in China by Long (1982), Zhang and Chen (1984), Wang et al. (1995);
and in India by Sharma (1969; G. microcos = M. paniculata).
Among the Peninsular Malaysian and Bornean Microcos species examined, three
main groups can be discerned based on whether the pollen is prolate-spheroidal,
248 Gard. Bull. Singapore 55 (2003)
16’ m
Plate 1. LMs and SEMs of selected Grewia and Microcos species showing tricolporate grains in polar
view. A & C: G. laevigata (A: Whitmore 3729; C: Hashim SAN 33470). B: G. multiflora (Motley 240).
D: M. fibrocarpa (Kochummen 80640). E: M. paniculata (Cramer 3405). F: M. malayana (Loh FRI
19249). Scale bar = 1 um in F; 10 um in A-E.
Pollen morphology of Grewia and Microcos 249
Plate 2. LMs and SEMs of selected Grewia and Microcos species showing tricolporate grains in equatorial
view. A: G. laevigata (Whitmore 3729). B: G. laevigata (Hashim SAN 33470) with lolongate endoaperture.
C: G. multiflora (Motley 240) with lalongate endoaperture. D: M. fibrocarpa (Kochummen 80640). E:
M. dulitensis (Richards 1767) with circular endoaperture. F: M. riparia (Hallier 2208) with circular
endoaperture. Scale bar = 1 1m in F; 10 um in A-E.
250 Gard. Bull. Singapore 55 (2003)
Plate 3. SEMs of selected Grewia and Microcos species showing exine detail of coarsely reticulate
pattern (A—D) or finely reticulate pattern (E—H) and lumina detail. A: G. multiflora (Motley 240), muri
variable in width and shape. B: G. laevigata (Hashim SAN 33470), muri branching inwards toward the
lumina forming reticulate subrugulate sculpturing. C: G. polygama (Kiah SFN 35286), lumina occupied
by a few microreticulations/perforations. D: G. huluperakensis (Turner 94-94), lumina occupied by
numerous microreticulations. E: M. henrici ssp. acuta (Ilias § 45121), muri variable in width and shape.
F: M. kinabaluensis (Chew & Corner RSNB 4994). G: M. dulitensis (Richards 1767), lumina occupied
by a few microreticulations. H: M. membranifolia (Singh SAN 30672), lumina occupied by numerous
microreticulations. Scale bar = 1 um in A-H.
Pollen morphology of Grewia and Microcos 251
Plate 4. TEMs micrograph of selected Grewia and Microcos species showing structure of ectexine and
endexine. A: G. occidentalis (Moss 3959). B: G. multiflora (Motley 240). C: M. paniculata (Kanodia
88480). D: M. tomentosa (Suppiah FRI 28239). Scale bar = 0.36 um in D; 0.45 um inA & C; 0.59 um
in B. Abbreviation used: c = columellae; cp = cytoplasm; ed = endexine; fl = foot layer; i = intine; sc =
spaces between columellae; t = tectum.
subprolate or prolate. In species with subprolate pollen grains, three subgroups can
be distinguished based on whether the endoapertures are circular, lalongate or circular-
lalongate. Other pollen morphological characters, e.g. P/E ratio and number of lumina
perforations, can be used to further subdivide these subgroups. Likewise, in species
with prolate pollen grains, by using palynological characters such as the number of
perforations of the lumina, P/E ratios and endoapertures shape, further subdivisions
to smaller groups can be made (Table 3).
Discussion
The pollen morphology of both genera is very similar and adds further confirmation
that species of Grewia and Microcos in Peninsular Malaysia and Borneo have pollen
morphological characters belonging to the ‘Grewia-type’ of Erdtman (1952).
Gard. Bull. Singapore 55 (2003)
252
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Pollen morphology of Grewia and Microcos 253
On the basis of palynological characters, the genus Microcos in Peninsular
Malaysia and Borneo can be distinguished from Grewia by a combination of the
following characters: pollen size, exine thickness, tectum ornamentation and lumina
diameter. The main morphological differences between pollen of Grewia and Microcos
are summarised in Table 4. To some extent, this study supports Burret’s view (1926)
for segregating Microcos from Grewia.
Table 4. Main differences between pollen of Grewia and Microcos species in Peninsular
Malaysia and Borneo.
Genus: Grewia Microcos
Characters
Size
Length polar axis, P (um) (44—)53-—75(-80) (20—)25-—35(-42)
Equatorial diameter, E (um) (32—)39-55(—62) (15—)19-27(-31)
Exine
Thickness (Um) (1—)1.55—2.15(—2.5) (0.70)0.84—0.90(-1)
Tectum
Ornamentation coarsely reticulate finely reticulate
Lumina diameter (um) = 13 at
Chung (2001, 2002) identified a total of 42 morphological and micromorphological
characters, which in combination can be used to distinguish Grewia from Microcos
species in Peninsular Malaysia and Borneo, and concluded that the two genera are
distinct. The findings support previous conclusions reached by Burret (1926) and
Bayer and Kubitzki (2003) based on the morphology of inflorescences, flowers and
fruits; by Chattaway (1934) based on wood anatomy; Zhang and Chen (1984) based
on pollen morphology; and by Bayer et al. (1999) based on combined analyses of
plastid atpB and rbcL DNA sequences.
Conclusions
This study confirms that within the Tiliaceae the pollen of Grewia and Microcos
species occurring in Peninsular Malaysia and Borneo conforms well with the so-
called ‘Grewia-type’ (Erdtman, 1952). Comparison based on results of this study and
all available published information indicates that pollen of Microcos differs from that
of Grewia in having smaller size, thinner exine, finer tectum reticulation and smaller
lumina, which supports the segregation of Microcos from Grewia.
254 Gard. Bull. Singapore 55 (2003)
Acknowledgements
The first author thanks the Director-General of the Forest Research Institute Malaysia
for the support and encouragement; we thank the Directors/Keepers/Curators of BM,
BO, BRUN, K, KEP, KLU, L, SAN, SAR and SING for the loan of specimens used
in this study; Professor Siwert Nilsson (Swedish Museum of Natural History) and Dr
R. W. J. M. van der Ham (L) for providing references; Mr Roslee Halip and Mr T.
Ragunathan for technical assistance; and Dr Madeline Harley (K) and Dr Ruth Kiew
(SING) for constructive comments that greatly improved the manuscript. Financial
support to the first author from the Tree Flora of Sabah and Sarawak Project IRPA
grant no. 01-04-01-0024 and the University of Malaya postgraduate research grant
nos. 489/97, 225/98, and 209/99 is gratefully acknowledged.
References
Bayer, C.,M.F. Fay, A. Y. de Bruijn, V. Savolainen, C. M. Morton, K. Kubitzki, W.S.
Alverson and M. W. Chase. 1999. Support for an expanded family concept of
Malvaceae within a recircumscribed order Malvales: a combined analysis of plastid
atpB and rbcL DNA sequences. Botanical Journal of the Linnean Society 129:
267-303.
Bayer, C. and K. Kubitzki. 2003. Malvaceae. Pp. 225-311. In: K. Kubitzki and C.
Bayer (eds.), The Families and Genera of Vascular Plants, Vol. 5 Flowering
Plants Dicotyledons. Springer-Verlag, Berlin.
Burret, M. 1926. Beitrage zur kenntnis der Tiliaceae I. Notzblatt des Botanischen
Gartens und Museum zu Berlin-Dahlem 9: 592-880.
Chattaway, M. M. 1934. Anatomical evidence that Grewia and Microcos are distinct
genera. Tropical Woods 38: 9-11.
Chaudhuri, S. K. 1965. Pollen morphological studies of the order Malvales II. Bulletin
of the Botanical Society of Bengal 19: 147-158.
Chaudhuri, S. K. and N. Mallik. 1965. Pollen morphological studies of the order
Malvales I. Bulletin of the Botanical Society of Bengal 19: 32-36.
Christensen, P. B. 1986. Pollen morphology studies in the Malvaceae. Grana 25: 95—
117.
Pollen morphology of Grewia and Microcos 255
Chung, R. C.K. 2001. Taxonomic and Micromorphological Studies of Grewia L. and
Microcos L. (Tiliaceae) in Peninsular Malaysia and Borneo. Unpublished Ph.D
Thesis, University of Malaya, Kuala Lumpur. 563 pp.
Chung, R. C. K. 2002. Leaf epidermal micromorphology of Grewia L. and Microcos
L. (Tiliaceae) in Peninsular Malaysia and Borneo. Gardens’ Bulletin Singapore
54: 263-286.
Coetzee, J. and H. P. Van Der Schijff. 1978. Pollen morphology of South Africa
Malvales: 1. Characteristics useful for keying and for numerical analysis. Journal
of South Africa Botany 45: 93-126.
El Ghazali,G. E. B. 1993. A study on the pollen flora of Sudan. Review of Palaeobotany
and Palynolology 76: 99-345.
Erdtman, G. 1952. Pollen Morphology and Plant Taxonomy: Angiosperms (An
introduction to palynology I). Almqvist & Wiksell, Stockholm. 559 pp.
Erdtman, G. 1960. The acetolysis method. A revised description. Svensk Botanisk
Tidskrift 54: 561-564.
Long, H. 1982. Tiliaceae. In: Angiosperm Pollen Flora of Tropic and Subtropic China.
Institute of Botany and South China Institute of Botany, Academia Sinica. The
Science Press, Beijing. Pp. 377-380 + plates 180-182.
Martinez-Hernandez, E., P. Fernandez and S. Lozano. 1978. Pollen of tropical trees.
I. Tiliaceae. Journal of the Arnold Arboretum 59: 299-309.
Nair, P. K. K. 1962. Pollen grains of Indian Plants HI. Bulletin of the Natural Botany
Garden 63: 1-33.
Nilsson, S. and A. Robyns. 1986. Bombacaceae. World Pollen and Spore Flora 14:
1-59.
Punt, W.,S. Blackmore, S. Nilsson and A. L. Thomas. 1994. Glossary of Pollen and
Spore Terminology. LPP Contribution Series No. 1. LPP Foundation, Utrecht. 71
PP:
Reynold, E.S. 1963. The use of lead citrate at high pH as electron opaque stain in
electron microscopy. Journal of Cell Biology 17: 208.
256 Gard. Bull. Singapore 55 (2003)
Sharma, B. D. 1969. Pollen morphology of Tiliaceae in relation to plant taxonomy.
Journal of Palynology 5: 7-29.
Sudhakar, S. and R.S. Rao. 1987. Biosystematic studies in the genus Grewia Linn. I.
Palynology. Journal of the Economic Taxonomy Botany 10(2): 437-441.
Wang, F.S., N. F. Chen, Y. L. Zhang and H. Q. Yang. 1995. Pollen Flora of China.
2nd edition. Institute of Botany, Academia Sinica, Beijing. Pp. 411-415, plates
194-196.
Zhang, Y. L. and Y. S. Chen. 1984. Studies on pollen morphology in Tiliaceae of
China. Acta Phytotaxonomica Sinica 22: 367-377.
Gardens’ Bulletin Singapore 55 (2003) 257-270
Six New Species of Bulbophyllum (Orchidaceae)
from Sulawesi
J.J. VERMEULEN
Singapore Botanic Gardens
Singapore 259569
AND
P.O’BYRNE
14 Clementi Crescent
Singapore 599517
Abstract
Six new species of Bulbophyllum are described, all from Sulawesi: B. trichorhachis (sect.
Hybochilus), B. valeryi (sect. Leptopus), B. auriculatum and B. tubilabium (both sect.
Monilibulbus), B. stenurum (sect. Osyricera), and B. reductum (sect. Pelma).
Introduction
The six new species described below once more underline the uniqueness of the
Sulawesi orchid flora. The dual origins of its flora are demonstrated by the fact that,
for instance, Bulbophyllum reductum belongs to an almost exclusively Papuan section
of the genus, whereas B. auriculatum and B. tubilabrum have their phylogenetic roots
in Asia, particularly the Sunda Shelf. B. stenurum, together with B. hastiferum Schltr.,
and B. anakbaruppui J.J. Verm. & P. O’Byrne, form a small, very distinctive series
within sect. Osyricera entirely restricted to Sulawesi; they demonstrate the amount
of speciation that has occurred on the island. B. trichorhachis is the second Sulawesi
species of sect. Hybochilus with a racemose inflorescence. This character is otherwise
almost absent in sect. Hybochilus as well as in assumedly related sections. Clearly,
the Sulawesi flora deserves more attention than it gets at present.
Bulbophyllum sect. Hybochilus
Bulbophyllum trichorhachis J.J. Verm. & P. O’Byrne, sp. nov. — Fig. 1.
258 Gard. Bull. Singapore 55 (2003)
Bulbophyllum trichorhachis J.J. Verm. & P.O’ Byrne, a Bulbophyllo iteranti floribus
multo minoribus (sepalum medianum 3.8 mm longum), labello carina
medialis carenti differt. —TYPE: Indonesia, Sulawesi, central part, SBG-
04348 (SING, holo.).
Roots below the pseudobulbs. Rhizome creeping, 0.8—1 mm diam., sections
between pseudobulbs 0.2—0.6 cm long, bracts not persistent. Pseudobulbs close
together, ovoid, 0.4—0.65 x 0.2-0.4 cm. Petiole up to 0.02 cm long. Leaf blade elliptic
to (ob)ovate, 1 .3-1.9 x 0.35—-0.5 cm, index (length/width) 2.6—5.4; acute. Inflorescence
an elongated raceme, 3-4.8 cm long, 1—22-flowered. Peduncle erect to patent, 2.5—2.9 cm,
bracts 3, the longest c. 1.5 mm long. Rhachis porrect or arching, zigzag bent, not
thickened, up to 1.7 cm. Floral bracts tubular at the base, 0.7—1 mm, acute. Flowers
not fully opening, each one developing only after anthesis of the previous. Pedicel
and ovary |.9-2.8 mm long, basal node on ac. 0.7 mm-long stump. Median sepal
recurved, ovate,c.4 x 1.5 mm, index 2.6—2.7; subacute, margins entire, base broadly
attached; thin, 3-veined, glabrous. Lateral sepals 4 x 1.5 mm, index 2.6—2.7; lower
margin somewhat ciliolate; otherwise as the median sepal. Petals porrect, ovate, c.
2.9 x 1.2 mm, index 2.4—2.5; subacute, margins ciliate, base narrowly attached; thin,
1-veined, glabrous. Lip slightly recurved near the base, general outline triangular, c.
2.9 x 0.7 mm, index c. 4.1 (all without artificial spreading); rounded, margins ciliate;
rather thick; adaxially concave near the base, convex elsewhere, surface glabrous;
abaxially without a ridge, surface with stiff hairs near the base, and with softer hairs
elsewhere. Column c.0.9 mm long, stigma without a tooth at its base, column foot
absent. Stelidia porrect, triangular, c.0.6 mm long, acute, with a slight, rounded wing
along the upper margin. Anther abaxially with a distinct, papillose crest, surface
otherwise finely papillose, front margin drawn out into an incurved, truncate beak
with entire margins. Pollinia 4; ovoid, the inner c. 3/4 as long as the outer; no
appendages present.
Colour: Flowers entirely dark purple.
Habitat: Epiphyte in montane forest, on branches of medium-sized trees. Alt.
c. 1000 m.
Distribution: INDONESIA: Sulawesi, central part (1 specimen seen).
Notes: Similar to Bulbophyllum iterans J.J. Verm & P. O’Byrne, differs in
having an unthickened, more elongated rhachis, smaller flowers (median sepal in
B. iterans\10.5—11.2 mm long, in B. trichorhachis c.4. mm long) and a lip without a
median ridge.
Figure 1. Bulbophyllum trichorhachis J.J. Verm. & P. O’Byrne — a. Habit; b. Inflorescence; c. Flower
analysis, from left to right: median sepal, petal, lateral sepal, lip; d. Lip, left: adaxial side, right: abaxial
side; e. Column and lip, lateral view; f. Anther, above: adaxial side, below: abaxial side; g. Pollinia,
above: two pairs, below: a single pair. — All from SBG-O 4348 (spirit sample).
Bulbophyllum from Sulawesi 259
260 Gard. Bull. Singapore 55 (2003)
Bulbophyllum sect. Leptopus
Bulbophyllum valeryi J.J. Verm. & P. O’Byrne, sp. nov. — Fig. 2.
Bulbophyllum valeryi J.J. Verm. & P.O’ Byrne, a Bulbophyllo melinantho pedunculo
2 mm longo, infra stigmatem umbonibus duobus distinctis differt. —
TYPE: Indonesia, Sulawesi, central part, SBG-O 4834 (SING, holo.).
Roots below the pseudobulbs. Rhizome creeping, 1-1.5 mm diam., sections
between pseudobulbs 0.25—0.4 cm long, bracts not persistent. Pseudobulbs close
together, ovoid, 0.45—0.7 x 0.3-0.55 cm. Petiole up to 0.1 cm long. Leaf blade elliptic
to ovate, 1.2—2.8 x 0.4—0.5 cm, index (length/width) 3—5.6; obtuse to acute.
Inflorescence \—1.2 cm long, 1-flowered. Peduncle patent, 0.15—0.2 cm, bracts 2, the
longest c. 1.8 mm long. Floral bract tubular, c.2 mm, acute. Flowers not fully opening.
Pedicel and ovary c.4 mm long, basal node on ac. 1.2 mm-long stump. Median sepal
recurved, triangular, c. 5.3 x 2 mm, index 2.6—2.7; acute, margins slightly and very
finely papillose, base widely attached; thick, glabrous. Lateral sepals free, oblique, c.
5.8 x 2.2 mm, index 2.6—2.7; margins glabrous, otherwise as the median sepal. Petals
porrect, ovate,c.3.3 x 2mm, index | .6—1.7; acute, margins slightly erose and papillose,
base narrowly attached; thin, glabrous. Lip hardly recurved, ovate, c. 2.8 x 1.2 mm,
index 2.3—2.4 (all without artificial spreading), obtuse, margins papillose; thick;
adaxially concave and with a median furrow in the basal half, slightly convex in the
top half, surface glabrous but papillose towards the margins about half-way up the lip
and with a patch of scattered glandular hairs in the top half; abaxially with a retuse
median ridge near the base, surface finely papillose, coarser papillose along the margins
towards the tip. Column c. 1.2 mm long, stigma with 2 distinct, papillose knobs at its
base, column foot with retrorse, triangular, obtuse wings near the tip. Stelidia triangular,
c.0.5 mm long, acute. Anther abaxially with a slight, rounded crest, surface finely
papillose, front margin not drawn out. Pollinia 4; ellipsoid, the inner c. as long as the
outer; no appendages present.
Colour: Sepals translucent pale ateentae suffused with ochrish red, veins
ochrish red. Petals translucent, some dark purple specks near the base, tip entirely
dark purple. Lip white, heavily suffused with purple.
Habitat: Epiphyte growing in small clusters on thin branches in montane
forest. Alt. c. 1000 m.
Distribution: INDONESIA: Sulawesi, central part (1 specimen seen).
Figure 2. Bulbophyllum valeryi J.J. Verm. & P.O’ Byrne — a. Habit; b. Flower; c. Flower analysis, from
left to right: median sepal, petal, lateral sepal, lip; d. Lip, left: adaxial side, right: abaxial side; e. Column
and lip, lateral view; f. Anther, left: adaxial side, right: abaxial side; g. Pollinia, left: two pairs, right: a
single pair. — All from SBG-O 4834 (spirit sample).
261
Bulbophyllum from Sulawesi
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262 Gard. Bull. Singapore 55 (2003)
Notes: This belongs to a suite of species within sect. Leptopus that is
characterized by densely clustered pseudobulbs on a creeping or shortly ascendent
rhizome, and a lip shape approaching what is commonly found in sect. Fruticicola
and in sect. Epibulbon. The species most similar to Bulbophyllum valeryi is B.
melinanthum Schltr, from New Guinea. B. valeryi differs in having a very short
peduncle and two distinct knobs below the stigma.
Named after Valeryus, son of Joseph Nasaret, who discovered this species.
Bulbophyllum sect. Monilibulbus
Bulbophyllum auriculatum J.J. Verm. & P. O’Byrne, sp. nov. — Fig. 3.
Bulbophyllum auriculatum J.J. Verm. & P.O’ Byrne, a Bulbophyllo minutulo et B.
schefferi ad labelli basin lobis lateralibus antrorsis triangularibus obtusis
differt. — TYPE: Indonesia, Sulawesi, SBG-O 4360 SING, holo.).
Roots below the pseudobulbs. Rhizome creeping, |.2—1.5 mm diam., sections between
pseudobulbs 0.25—0.6 cm long, bracts not persistent. Pseudobulbs orbicular to narrowly
ovoid, basal half prostrate on and fused to the rhizome, so that the new pseudobulbs
arise c. half-way up the old, 0.35-0.9 x 0.28-0.35 cm. Petiole 1.8—3 mm long. Leaf
blade ovate to elliptic, 0.65—1 x 0.22—0.32 cm, index (length/width) 2—3.5; acuminate.
Inflorescence c. 5 cm long, 1-flowered. Peduncle erect to patent, c. 2.5 cm, bracts 2,
the longest 3—3.5 mm long. Floral bracts tubular, c. 2 mm, acute. Flowers not fully
opening. Pedicel and ovary c. 16.5 mm long, basal node on ac. 1.4 mm-long stump.
Median sepal porrect with recurved top, ovate, c. 6.2 x 1.8 mm, index 3.43.5; acute,
margins slightly erose-papillose towards the tip, base rather widely attached; thin,
glabrous. Lateral sepals recurved, free, oblique, triangular, c. 7 x 2.3 mm, index 3—
3.1; otherwise as the median sepal. Petals porrect, ovate, c. 2.3 x 0.8 mm, index c.
2.9; acute, margins slightly erose-papillose towards the tip, base widely attached;
thin, glabrous. Lip recurved at c. 1/3 of its length, + 3-lobed; midlobe ovate, c.2.4 x
1.2 mm, index c. 2 (all without artificial spreading), rounded, margins very finely
papillose; thick; adaxially concave and with 2 short ridges near the base converging
distally and then fused to a small callus, beyond this callus a median furrow reaching
up to about half-way the lip, adaxial surface papillose in the centre; abaxially with
barely a median ridge, top part slightly concave, surface glabrous; lateral lobes erect,
antrorse, attached along the proximal third of the lip, obliquely triangular, c.0.5 mm
high, obtuse, thin, glabrous. Column c. 1.4 mm long, rostellum thick, knob-shaped,
Figure 3. Bulbophyllum auriculatum J.J. Verm. & P. O’Byrne — a. Habit; b. Flower; c. Flower
analysis, from left to right: median sepal, petal, lateral sepal, lip; d. Lip, left: adaxial side,
right: abaxial side; e. Column and lip, lateral view; f. Anther, left: adaxial side, right: abaxial
side. — All from SBG-O 4360 (spirit sample).
263
Bulbophyllum from Sulawesi
264 Gard. Bull. Singapore 55 (2003)
stigma without a tooth at its base, column foot without teeth. Srelidia triangular, c.0.4
mm long, acute, with a small, antrorse, obliquely deltoid, rounded to obtuse tooth
along the upper margin. Anther abaxially with a widely rounded crest, surface +
glabrous, front margin somewhat drawn out into a triangular, obtuse beak. Pollinia:
not seen.
Colour: Sepals yellowish orange. Lip red.
Habitat: Understorey epiphyte in montane forest, at c. 1800 m alt. Flowering
observed in Sept.
Distribution: INDONESIA. Sulawesi, central part (1 specimen seen).
Notes: Most similar to Bulbophyllum minutulum Ridl., and B. schefferi (O.
Kuntze) Schltr., Bulbophyllum auriculatum differs from both in having distinct lateral
lobes at the base of the lip.
Bulbophyllum tubilabrum J.J. Verm. & P. O’Byrne, sp. nov. — Fig. 4.
Bulbophyllum tubilabrum J.J. Verm. & P. O’Byrne, a Bulbophyllo hodgsonii sepalo
mediano brevi lato (index 1.7—1.8 contra 4—5 in B. hodgsonii), floribus
multo minoribus (sepala lateralia c. 4.8 mm longa contra c. 15 mm in B.
hodgsonit) differt.— TYPE: Indonesia, Sulawesi, SBG-O 4935 (SING, holo.).
Peduncle erect to patent, c. 2.5 cm, bracts 2, the longest c. 2 mm long. Floral
bracts tubular, c. 1.7—2 mm, acute. Flowers fully opening. Pedicel and ovary c. 10
mm long, basal node on a c. 1.2 mm-long stump. Median sepal porrect, ovate-
triangular,c.3 x 1.7 mm, index |.7—1.8; acute-acuminate, margins entire, base widely
attached; thin, glabrous. Lateral sepals recurved, free, oblique, elliptic-ovate, c.4.8 x
2.8 mm, index c. 1.7; acute; otherwise as the median sepal. Petals porrect, ovate, c.
1.4x 0.9 mm, index | .5—1.6; subacute, margins entire, base rather narrowly attached;
thin, glabrous. Lip somewhat recurved at c. 1/4 of its length, triangular near the base,
cylindrical elsewhere with the margins folded back over the abaxial side and fused, c.
3.4 x 1.5 mm, index c. 2.2—2.3 (all without artificial spreading), rounded, margins
entire; thick, glabrous; adaxially concave near the base, with a small cavity just above
the ligament, ending in a knob that distally continues as a median strip of slightly
thickened tissue ending about half-way up the lip as a drop-shaped callus, and with 2
short ridges bordering this median strip from 1/5 to 2/5 of the lip, abaxially with a
rounded median ridge near the base. Column c. 1 mm long, stigma without a tooth at
its base, column foot without teeth. Stelidia triangular, c. 0.4 mm long, acute, with a
small, deltoid, rounded tooth along the lower margin. Anther abaxially with a widely
rounded crest, surface papillose, front margin not drawn out. Pollinia: not seen.
Figure 4. Bulbophyllum tubilabrum J.J. Verm. & P.O’ Byrne — a. Flower; b. Flower analysis, from left to
right: median sepal, petal, lateral sepal, lip; c. Lip, above: adaxial side, below: abaxial side; e. Column
and lip, lateral view; f. Anther, above: adaxial side, below: abaxial side. — All from SBG-O 4935 (spirit
sample).
265
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266 Gard. Bull. Singapore 55 (2003)
Colour: Sepals yellowish. Lip orange red.
Habitat: Understorey epiphyte in montane forest, at c. 1800 m alt. Flowering
observed in Sept.
Distribution: INDONESIA. Sulawesi, central part (1 specimen seen).
Notes: Bulbophyllum hodgsonii Hend. is most similar but has larger flowers
with narrower sepals: median sepal 12-15 x 3 mm, index 4-5, lateral sepals c. 15 mm
long.
Bulbophyllum sect. Osyricera (= sect. Globiceps)
Bulbophyllum stenurum J.J. Verm. & P. O’Byrne, sp. nov. — Fig. 5.
Bulbophyllum stenurum J.J. Verm. & P. O’Byrne, a Bulbophyllo hastifero rhachidi
non incrassata differt. — TYPE: Indonesia, Sulawesi, central part, SBG-O
4902 (SING, holo.).
Roots creeping along the rhizome towards its point of attachment, then
spreading over the substrate. Rhizome + pendulous with + patent branches, up to 50
cm long, 4.5—5 mm diam., sections between pseudobulbs 3—5.6 cm long, bracts
persistent. Pseudobulbs distant, ovate, 3.6—-4.8 x 1—1.2 cm, distinctly and rather sharply
4—5-angled. Petiole c. 1 cm long. Leaf blade ovate, c. 19 x 2.2 cm, index (length/
width) 8.6—-8.7; acute. Inflorescence porrect to recurved, a rather dense raceme, c. 6.5
cm long, c. 15-flowered. Peduncle c. 1.4 cm, bracts 2, the longest c. 8 mm long.
Rhachis not thickened, c. 5 cm long. Floral bracts c. 3.5 mm, acute. Flowers not
resupinate, spirally arranged, not fully opening, many open simultaneously. Pedicel
and ovary 2.4—2.7 mm long, basal node + flush with the surface of the rhachis.
Median sepal + porrect, ovate,c.4.5 x 3.1 mm, index 1.4—1.5; top cucullate, rounded,
margins entire, papillose-ciliolate, base rather narrowly attached; thin, 5-veined, surface
adaxially glabrous, abaxially shortly hirsute. Lateral sepals adnate along the lower
margins, basal part recurved, top part incurved, oblique, triangular, c. 5.6 x 2.4mm,
index 2.32.4, top flat, subacute, base broadly attached; 3-veined, otherwise as the
median sepal. Petals recurved, ovate-triangular, c. 4 x 0.9 mm, index 4.44.5; acute,
margins entire, ciliolate, base narrowly attached; very thin, 1-veined, surface adaxially
finely papillose towards the tip, abaxially glabrous. Lip about straight, general outline
ovate, c. 4 x 1.7 mm, index 2.3—2.4 (all without artificial spreading, hastate with a
drawn out, elliptic midlobe and deltoid, acute sidelobes when spread); obtuse, margins
entire; rather thick, glabrous; adaxially slightly concave near the base, with 2 parallel,
Figure 5. Bulbophyllum stenurum J.J. Verm. & P.O’ Byrne — a. Habit; b. Flower; c. Flower analysis,
from left to right: median sepal, petal, lateral sepal, lip; d. Lip, left: adaxial side, right: abaxial side; e.
Column and lip, lateral view; f. Anther, left: adaxial side, right: abaxial side; g. Pollinia, left: two pairs,
right: a single pair. — All from SBG-O 4902 (spirit sample).
Bulbophyllum from Sulawesi
267
Mem eee ttre eeeee®
268 Gard. Bull. Singapore 55 (2003)
short, inconspicuous, rounded ridges at c. 1/3 of the length of the lip, adaxial surface
glabrous; abaxially + without a ridge near the base, surface papillose in the distal 1/3
of the lip. Column c. 2 mm long, stigma without a tooth at its base, column foot with
2 truncate wings near its base, and a conspicuous swelling just above the ligament.
Stelidia minute, abruptly bent downwards but with an antrorse, + acute tip, along the
upper margin with a very large (far more conspicuous than the stelidia themselves),
antrorse, wedge-shaped wing with a broadly rounded, erose top margin. Anther
abaxially with a crest largely overtopping the front margin, surface coarsely papillose,
front margin not drawn out. Pollinia 4, the inner + as long as the outer, all ellipsoid-
ovoid; no appendages present.
Colour: Unknown.
Habitat: Epiphyte in montane forest. Alt. c. 1000 m.
Distribution: INDONESIA: Sulawesi, central part (1 specimen seen).
Notes: Bulbophyllum hastiferum Schltr. is most similar but differs in having
a thickened, spindle-shaped rhachis. it also has a narrower median sepal, a lip with a
narrower midlobe, and an anther with a wider abaxial crest.
Bulbophyllum sect. Pelma
Bulbophyllum reductum J.J. Verm. & P. O’ Byrne, sp. nov. — Fig. 6.
Bulbophyllum reductum J.J. Verm. & P. O’Byrne, a Bulbophyllo savaiensi ssp.
subcubico columnae pede carenti, ad labelli basin sine cavitate distincta differt.
— TYPE: Indonesia, Sulawesi, central part, SBG-O 3886 (SING, holo).
Roots growing along the rhizome towards its point of attachment, then spreading
over the substrate. Rhizome erect to patent, up to 10 cm long, 1.8—2.4 mm diam.,
sections between pseudobulbs 0.8—1.2 cm long, bracts persistent. Pseudobulbs distant,
ovoid, 1.2—1.6 x 0.25—0.4 cm. Petiole 0.05—0.1 cm long. Leaf blade ovate, 2.3-3.4 x
0.75—0.85 cm, index (length/width) 3-4; acute. Inflorescences along the entire rhizome,
porrect or recurved, lax racemes, 0.5—1.05 cm long, 1—4-flowered. Peduncle 0.3—
0.5 cm, bracts 4—5, the longest 2—3.5 mm long. Rhachis up to 0.55 cm. Floral bracts
tubular proximally, 1 .4—1.9 mm long, acuminate. Flowers not resupinate, little opening,
several open simultaneously. Pedicel and ovary | .3—1.7 mm long, basal node on ac.
0.6 mm-long stump. Median sepal porrect, ovate, c. 1.4 x 0.8 mm, index 1.7—1.8;
rounded and apiculate, margins papillose towards the tip, base broadly attached; thick,
Figure 6. Bulbophyllum reductum J.J. Verm. & P. O’Byrne — a. Habit; b. Inflorescence; c. Flower
analysis, from left to right: median sepal, petal, lateral sepal, lip; d. Lip, left: adaxial side, right: abaxial
side; e. Column and lip, lateral view; f. Anther, above: adaxial side, below: abaxial side; g. Pollinia,
above: two pairs, below: a single pair. — All from SBG-O 3886 (spirit sample).
269
Bulbophyllum from Sulawesi
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270 Gard. Bull. Singapore 55 (2003)
1-veined, glabrous. Lateral sepals ovate-triangular, c. 1.7 x 0.8 mm, index 2.1—2.2;
acuminate, margins entire; otherwise as the median sepal. Petals porrect, obovate, c.
0.9 x 0.55 mm, index 1.6—1.7; truncate-rounded, margins papillose towards the tip,
base rather narrowly attached; thin, l1-veined, glabrous. Lip slightly recurved at c.
3/5 of its length, general outline elliptic, c. 0.8 x 0.7 mm, index 1.1—1.2 (all without
artificial spreading); truncate-rounded, margins entire; rather thick, glabrous; adaxially
slightly concave near the base, with a slight transverse callus about half way, top half
concave with a transverse, triangular, subacute, thick tooth in the centre; abaxially
without a ridge. Column c. 0.7 mm long, stigma without a tooth at its base, column
foot absent. Stelidia about absent. Anther abaxially without a crest, surface papillose,
front margin not drawn out. Pollinia 4; ovoid, the inner c. 2/3 as long as the outer; no
appendages present.
Colour: Flowers cream-coloured.
Habitat: Epiphyte in mossy montane forest. Alt. 1100-1200 m.
Distribution: INDONESIA: Sulawesi, central part (1 specimen seen).
Notes: Deceptively similar in general habit to Bulbophyllum savaiense Schltr.
ssp. subcubicum (J.J. Sm.) J.J. Verm. Only when analyzing the minute flowers can B.
reductum be distinguished at once by the absence of a column foot, and of a distinct
cavity at the base of the lip. The absent column foot makes inclusion of B. reductum
in sect. Pelma problematic, but the marked overall similarity between B. reductum
and B. savaiense leaves no other option.
Gardens’ Bulletin Singapore 55 (2003) 271— 307
A Botanical Survey of Chek Jawa,
Pulau Ubin, Singapore
SERENA LEE! , SAMSURI AHMAD’, PAUL LEONG’,
ALI IBRAHIM? AND AIK-TECK GWEE!
'Singapore Botanic Gardens, Singapore 259569
*Conservation Division, Operations Department,
National Parks Board,
Singapore 189720
Abstract
A botanical survey in 2002/2003 collected 245 species in 171 genera and 80 families of vascular
plants excluding alien species. Among the seven habitats on Chek Jawa, the intertidal zone
with seven species of seagrasses, the rocky shore and the mangrove communities are relatively
diverse and undisturbed. The coastal forest, although heavily disturbed, is not secondary forest.
Based on the Singapore Red Data Book, 11 species were recorded as ‘extinct’, 11 as endangered
and 16 as vulnerable. The re-finding of these extinct species makes Chek Jawa an important
conservation site in the Singapore context. Elephantopus mollis (Compositae) is a new record
for the Singapore flora.
Introduction
Pulau Ubin, an island situated northeast of Singapore, covers an area of 10.2 km?
measuring 1.5 km north to south and 8 km east to west. About a hundred residents
still live on the island (The Straits Times, 24 Aug 2002) and its rustic charm makes it
a popular outdoor attraction for city dwellers (Chua, 2000, 2003). Tanjung Chek Jawa
is a headland on the northeast tip of the island (Fig. 1) that includes the intertidal area
of about 100 ha and the hinterland of 10.5 ha (N 1° 0' 30" E 103° 59' 30" coordinates
at the site of beacon on top of the hill).
Chek Jawa became the focus of attention when its intertidal area was surveyed
and found to be extremely rich in marine life (Tan and Yeo, 2003). It was already
earmarked for land reclamation under the Development Guide Plan in 1997 and the
Master Plan in 1998 (URA, 2003). Public support for conserving Chek Jawa was
strong as it was one of the few areas of natural marine heritage left. Petitions to the
government to preserve the area were heeded and Chek Jawa was saved just nine
days prior to start of reclamation work (The Straits Times, 2 Jan 2002). The island
has been largely zoned as ‘open space and reserve land’ in the 2001 Concept Plan and
will now be left intact for at least 10 years (The Straits Times, 15 Jan 2002).
ade Gard. Bull. Singapore 55 (2003)
While attention by both scientists and the public has focused on the intertidal
ecosystem of Chek Jawa, the vegetation on the headland has never been subject to a
systematic survey. It was with this end in view that the survey was under taken to
provide a comprehensive checklist of vascular plants and to evaluate their conservation
status. Another reason for carrying out the survey was that there is very little coastal
forest left in Singapore and so it is important to document this habitat.
History of Botanical Collecting on Pulau Ubin
The earlist collections were made by Ridley, Hullet and Goodenough in the 1880s. A
few specimens were later collected by Furtado in the 1920s and Allen in the 1940s.
Those appear to be random samplings of the flora of Pulau Ubin. Neither the precise
locality of the collections nor their habitat was given. In the 1990s, Ali and Lai collected
15 and 20 specimens from Pulau Ubin, respectively. In 1990, Turner et al. (1992)
carried out a botanical survey of Pulau Ubin, of which two specimens were collected
from Chek Jawa. The online Checklist of Chek Jawa (Lai, undated; http://
habitatnews.nus.edu.sg/news/chekjawa/checklists.htm#1) was not supported by
herbarium specimens.
Methodology
Specimens were collected from October 2002 to October 2003 until no more new
records were found. However, about half of the area in the coastal forest habitat is
inaccessible and so was not surveyed due to the dense thickets of climbers, debris of
dead tree stumps and branches on the steep slope. Wherever possible, fertile material
was collected. In its absence, sterile material was also included. All specimens are
deposited and databased in the Herbarium, Singapore Botanic Gardens (SING).
Common weeds and alien species were collected but not included in the survey
Statistics.
Conservation status was assessed using the categories from The Singapore
Red Data Book (Ng & Wee, 1994) in which [Ex] denotes Extinct in Singapore; [En]
denotes Endangered; [V] denotes Vulnerable and [R] denotes Rare. In addition, [exotic]
denotes plants that do not originate in the region and [C] denotes Common. Long-
term viability of these plant populations is dependent on their ability to regenerate
and the security of their habitat.
A compilation of all previous collections in SING from Pulau Ubin was
compared with the current checklist.
Survey of Chek Jawa, Singapore
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Results and Discussion
Species Composition
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This survey collected a total of 245 species in 171 genera and 80 families of which 7
species were seagrasses, 20 were ferns, 3 were gymnosperms and 215 were terrestrial
flowering plants (Appendix). This compares with the survey carried out by Turner er
al. (1992) for the whole of Pulau Ubin, which listed 261 species with a further 71
274 Gard. Bull. Singapore 55 (2003)
species of aliens. This shows that Chek Jawa is an important site and a significant
component of the entire island of Ubin.
In Chek Jawa, the Leguminosae and Rubiaceae are the best represented
families (Table 1). The Leguminosae are represented by climbers and trees and includes
most of the smothering climbers, such as Aganope thyrsiflora, Derris amoena and
Entada spiralis. On the slope of the coastal forest, small trees and shrubs of the
Rubiaceae such as Aidia spp., Canthium spp. and Timonius spp. are common. The
common climber of this family, Morinda umbellata, is ubiquitous in the coastal forest:
on slopes, ridges as well as near the beach fringe. Moraceae are represented by species
of Artocarpus and Ficus, many of which are big trees. Many of the secondary forest
trees are represented by the Euphorbiaceae, such as species of Glochidion and
Macaranga. Annonaceae are represented by shrubs, such as Desmos chinensis, and
lianas, such as Artabotrys suaveolens and Uvaria spp. These are mainly found in the
coastal forest habitat.
Table 1. Comparison of the five most common plant families (excluding aliens)
collected in Chek Jawa and Pulau Ubin (Turner et al., 1992).
Chek Jawa No. of species Pulau Ubin No. of species
Leguminosae 18 Gramineae 18
Rubiaceae 17 Cyperaceae 18
Moraceae IZ Euphorbiaceae 16
Euphorbiaceae 11 Leguminosae 15
Annonaceae 2 Compositae 15
The most speciose families (Table 1) collected in the Pulau Ubin survey
(Turner et al., 1992) were grasses (Gramineae) and sedges (Cyperaceae). The high
representation of these families indicates that many of the collections were from the
grassland/wasteland habitats as does the inclusion of the Compositae. Secondary forest
species, such as species of Macaranga (Euphorbiaceae) and Mimosa (Leguminosae)
are also well represented. Leguminous climbers, such as Derris spp. Desmodium spp.
and Entada spiralis are plants that were commonly found both in the Pulau Ubin and
Chek Jawa surveys.
Elephantopus mollis (Compositae) is a new record for the Singapore flora.
Habitats
The sites covered in this survey included seven distinct habitats: the intertidal zone,
the rocky/sandy shores, mangrove, coastal forest, abandoned rubber plantation and
dusun (mixed fruit orchard) and patches of wasteland.
Survey of Chek Jawa, Singapore 275
Intertidal zone
Seagrass beds are becoming rare in Singapore. Tan and Yeo (2003) illustrated the
seagrasses found at Chek Jawa. We collected seven species of seagrass, excluding
Enhalus acoroides that is known to occur on the seaward side of the sandbar in water
too deep for collection at the time of survey. Three species belong to the Cymodoceaceae
and four to the Hydrocharitaceae. Out of these, three species, Halodule pinifolia,
Halodule uninervis (Cymodoceaceae) and Thalassia hemprichii (Hydrocharitaceae),
were listed as ‘extinct’: and two, Halophila beccarii and Halophila spinulosa
(Hydrocharitaceae), as endangered. The two most commonly found seagrasses were
Halophila ovalis and Halophila spinulosa (Hydrocharitaceae), both of which thrive
on sandbars with coral rubble. Also found on sandy areas were Halophila beccarii
and Halophila pinifolia, while Cymodocea rotundata, Halodule uninervis and
Thalassia hemprichii occurred on mud flats.
Rocky shore
Singapore has very few undisturbed rocky shore habitats. Plants around the rocky
shore face conditions similar to plants on the fringe of the coastal forest. Trees here
do not grow very tall; they are slow growing because of the adverse conditions and
the nutrient-poor soils. However, the height of some individual trees, such as
Memecylon edule (7 m), Garcinia hombroniana (8 m), Intsia bijuga (9 m), Pouteria
linggensis (10 m) can be considered exceptional by any standard. Garcinia
hombroniana is listed as an endangered species.
Mangroves
The mangrove pockets on Chek Jawa are rich in biodiversity, supporting 20 species
of mangrove plants, many of them mature, good-sized trees as well as 14 species of
transitional mangrove plants. Of the true mangrove plants, Bruguiera parvifolia (the
only one found in Chek Jawa) is endangered; Rhizophora stylosa and Xylocarpus
moluccensis are vulnerable. A Ceriops species was recently discovered, which is
substantially different from the adjacent C. tagal trees and has been re-identified as
C. decandra. Nypa fruticans, is found in brackish water. Transitional mangrove plants
were represented by trees such as Terminalia catappa, Cerbera manghas and Diospyros
ferrea; shrubs such as Allophylus cobbe, Clerodendrum inerme and Ximenia
americana; climbers such as Flagellaria indica, Salacia chinensis and Dalbergia
candenatensis; as well as the pandan, Pandanus odoratissimus and the palm,
Oncosperma tigillarium. Epiphytes such as Dischidia major, D. nummularia and Hoya
verticillata were common on transitional mangrove trees.
Coastal forest
The coastal forest is a unique habitat in that plants that inhabit the area are adapted to
276 Gard. Bull. Singapore 55 (2003)
the harsh conditions often associated with the coast. Strong, drying winds, salt spray,
intense light reflection from the sea and other factors can affect their growth and
survival. Trees along the edge of the hill cling precariously to the steep slope. They
were observed to grow outwards from the forest towards the sea. This could be the
result of simple phototropism or in some cases, soil erosion at the fringe of steep slope,
where a tree fell but was halted in its fall by surrounding vegetation and then continued
to grow with the main trunk at a sharp incline. Scaphium macropodum and Aidia
densiflora were examples of this.
Unlike the lowland dipterocarp forest, the coastal forest does not exhibit the
typical five-layer stratification. Coastal forest on Chek Jawa more resembles that
described by Corner (1985) for the flora of coastal islets on the east coast of Peninsular
Malaysia both in species composition and tree height (maximum 21 m).
On Chek Jawa, other than a few large trees, there is a lack of the emergent
layer. Below the low main canopy of trees (about 5 m tall) and climbers were sparse
populations of shrubs and treelets. On steep slopes, sparsely populated with trees,
their canopies struggle with the climbers for light. The largest trees observed (Table
2) were a Pouteria malaccensis tree about 7—8 m tall and a Chrysophyllum roxburghii
tree 8—10 m tall, the latter with three trunks, the largest, 0.5 m in diameter. These
taller trees were found on the more level part of the slope. Other trees observed growing
on the slope included Adenanthera malayana, Archidendron contortum and A.
ellipticum, Diospyros spp., Elaeocarpus pedunculatus, Eurycoma longifolia, Ficus
vasculosa, Garcinia atroviridis, G. hombroniana, Glochidion superbum,
Campylospermum serratum, Ixonanthes reticulata, Knema globularia, Neolitsea
zeylanica, Pouteria linggensis, P. obovata, Triadica cochinchinensis, Suregada
multiflora and Vitex pinnata. A clump of the monocot tree, Dracaena maingayi; the
palm Licuala spinosa and the clumping palm, Oncosperma tigillarium as well as the
‘extinct’ pandan Pandanus tetrodon were also observed. The ‘extinct’ climber, Gnetum
latifolium was also found here. The understorey layer was interspersed with shrubs
such as Clerodendrum laevifolium and saplings of Olea brachiata, Archidendron
contortum and other treelets. The flora, hence typifies the characteristics of a coastal
forest habitat. In alarming proportions are the climbers, some of which reached
substantial height and formed a dense mass that smothered the surrounding plants as
well as blocking sunlight from reaching the forest floor. Examples of these climbers
are Aganope thyrsiflora, Derris amoena, Entada spiralis, Linostoma pauciflorum
and Fibraurea tinctoria.
At the fringe of the coastal hill towards the landward side, on more level
terrain, where the forest merges with the abandoned rubber estate, plants of the more
typical secondary forest species were seen. Examples are climbers such as Embelia
ribes, Morinda umbellata and Tetracera indica; shrubs such as I[xora congesta,
Clerodendrum laevifolium and Memecylon amplexicaule (the last listed as vulnerable);
trees such as Fagraea racemosa, Syzygium lineatum, S. zeylanicum and Rhodamnia
Survey of Chek Jawa, Singapore aya
Table 2. The Tallest Trees at Chek Jawa
Height (m) Family Species Habitat
Zs Burseraceae Dacryodes rostrata Coastal Forest
20 Anacardiaceae Mangifera indica Coastal Forest
20 Bombacaceae Durio zibethinus Coastal Forest
18 Myristicaceae Knema globularia Coastal Forest
15 Dracaenaceae Dracaena maingayii — Coastal Forest
10 Meliaceae Xylocarpus granatum Mangrove
10 Moraceae Artocarpus dadah Coastal Forest
10 Moraceae Ficus kerkhovenii Coastal Forest
10 Moraceae Ficus superba Coastal Forest
10 Sapotaceae Chrysophyllum Coastal Forest
roxburghii
10 Sapotaceae Pouteria linggensis Rocky Shore
9 Leguminosae Intsia bijuga Rocky Shore
8 Apocynaceae Cerbera manghas Rocky Shore
8 Chrysobalanaceae Maranthes corymbosa Coastal Forest
8 Guttiferae Garcinia hombroniana_ Rocky Shore
8 Sonneratiaceae Sonneratia alba Mangrove
7-8 Sapotaceae Pouteria malaccensis Coastal Forest
cinerea were found. Of interest is the common occurrence of the ‘extinct’ tree Knema
globularia, albeit at the coastal fringes. The liana, Cnestis palala was found along the
fringe of a gravel pathway, some reaching 10 m high. Larger trees were mostly found
in this part of the forest where the forest floor is flatter and thus more stable (Table 2).
Abandoned rubber estate, dusun and wasteland
The area surrounding the coastal forest includes the long abandoned Ong Ting Lye
Rubber Estate and dusun (mixed fruit orchards) of the villages. Fruit trees that still
persist include Artocarpus altilis (breadfruit), A. heterophyllus (jackfruit), A. integer
(chempedak), Averrhoea carambola (starfruit), Durio zibethinus (durian), Garcinia
mangostana (mangosteen), Manilkara zapota (chiku), Nephelium lappaceum
(rambutan), Psidium guajava (guava), Spondias pinnata (kedondong) and Syzygium
aqueum (jambu ayer). These relics of cultivation are now all that remains of the people
who were re-located to the mainland. |
Escapes from cultivation, which include Tinospora crispa (a medicinal plant
used locally for malaria fever, jaundice and intestinal worms), Passiflora laurifolia
(passion fruit) and garden escapes, such as Gloriosa superba, still persist in these
areas.
278 Gard. Bull. Singapore 55 (2003)
Villagers and the plantation owner probably cleared tracks that are now
wasteland. Thickets of ferns, for example, Nephrolepis auriculata, Pityrogramma
calomelanos,Acrostichum aureum, and a number of sedges and grasses have invaded
these wasteland patches.
Plants of Interest
Enhalus acoroides
This seagrass species was found on the seaward side of a sandbar that is for most of
the time inundated by water. Its long leaves about 1.5 m are reported to be the food of
the dugong.
Fagraea racemosa
The big leathery leaves and inflorescences with peachy cream flowers are attractive
and could be used more frequently in horticultural planting.
Gnetum latifolum
This climber is listed as ‘extinct’, but was seen in various patches in the coastal
forest. Of particular interest, other than its status, is that it and G. microcarpum are
the only gymnosperms found growing wild here. (G. gnemon, meninjau, an escape
from cultivation, is also found on Chek Jawa). Cycas rumphii s.l. commonly found
on seashores is not found here.
Intsia bijuga
This valuable timber species can grow to an impressive height in more conducive
environments. Its pinkish white fragrant flowers and thick, round leaves are attractive.
Knema globularia.
The fruits of the sea nutmeg exhibit bright red pulp when ripe thus attracting larger
birds that eat and disperse the seeds. These trees are found on the seaward side of the
coastal forest and rocky shoreline and, because of its rarity elsewhere in Singapore,
has become an integral attraction of Chek Jawa.
Lasianthus hirsutus
This species, listed as ‘extinct’, is found along the edge of the gravel pathway at the
landward fringe of the coastal forest. Its large velvety leaves and stunning metallic
blue fruits lend it horticultural potential.
Memecylon edule
With its gregarious flowering and captivating blue-purple flowers, the plant has
become an icon of the rocky shoreline of Chek Jawa. When in flower, it adds a splash
Survey of Chek Jawa, Singapore 279
of blue-purple - a rare colour in the tropics, to the already picturesque scenery.
Large trees
The largest of all was Dacryodes rostrata with an impressive height of 25 m (Table 2)
and trunk | m in diameter. Along the rocky shore, a Knema globularia about 7 m tall
grew in a reclining position. In the mangroves, Xylocarpus granatum with its snake-
like roots anchored well into the substratum, provided support for its 10 m tall crown.
Orchids
Surprisingly, orchids were poorly represented. Only two common orchids were found.
The epiphytic Dendrobium crumenatum and the terrestial Eulophia graminea, the
latter growing on the sandy beach in the mangrove patch.
Hemiparasitic plants
Dendrophthoe pentandra was the only hemiparasite encountered in the survey. It was
seen perched on rubber trees about 10 m above the ground. It was too high to collect.
Disturbance
Turner et al. (1992) analysed the early collections from Pulau Ubin and concluded
that even in the 1880s, the vegetation was largely secondary with no primary forest
species. They went so far as to conclude that at one time the terrestrial vegetation
might have been cleared. The survey of Turner et al (1992) found that plant diversity
on Pulau Ubin was low and that the frequency of aliens was relatively high indicating
a high degree of human interference. Their species list comprised largely of common
weeds, secondary forest and mangrove species. Primary forest species were almost
completely lacking.
The coastal forest on Chek Jawa does not exhibit the species composition
typical of secondary forest. Corlett (1994) in discussing a definition of secondary
forest, mentioned that ‘virtually all tropical forests have suffered some form of human
impact’ and that ‘the question is: how much disturbance is needed to make a forest
secondary?’ In his opinion,‘the key difference is the break in the continuity of forest
occurrence on the site and the consequent dependence for recovery on dispersal from
outside.’ He mentioned that it would be preferable to use “primary’— qualified where
necessary by terms such as “logged”’, “depleted” or “degraded” — to indicate continuity
of forest presence at a site and to use “unmodified” or “virgin” to describe forests that
have not obviously been changed by significant human impact.
In Chek Jawa’s coastal forest, secondary forest species such as Adinanandra
dumosa, Ficus grossularioides, Macaranga spp. and Dillenia suffruticosa although
present, are not in abundance; whereas species such as Adenanthera malayana,
280 Gard. Bull. Singapore 55 (2003)
Archidendron contortum, A. ellipticum, Diospyros ferrea, Eurycoma longifolia,
Garcinia atroviridis, G. hombroniana and Knema globularia, which are not typical
representatives of secondary forest, occur there. The forest does show signs of heavy
disturbance but it is obvious that the area was not cleared in the past. Therefore, by
Corlett’s definition, the coastal forest of Chek Jawa is degraded primary coastal forest
not secondary forest. Further, 65 species listed by Corner (1985) on the uninhabited
and undisturbed coastal islets are also found in Chek Jawa.
The canopy, though, is not continuous or lofty. Most trees on the steep hill
are not very tall ranging between 4—6 m in height. This low canopy is indicative of a
history of disturbance. Part of the disturbance is undoubtably man-made as there are
remains of building foundations at the top of the hill. The bamboo (Bambusa vulgaris)
that occurs in clumps throughout Chek Jawa is one such indication of disturbance.
However, the frequency of trees near the slope in coastal forest with a decayed main
trunk and abundant coppicing, such as Chrysophyllum roxburghii that has side shoots
up to 0.5 m in diameter, suggests that in the past natural phenomena such as high
winds might have brought down the larger trees.
In contrast, the rocky shore vegetation does not show signs of disturbance
and many species are represented by large and presumably old specimens such as
Cerbera manghas, Garcinia hombroniana, Knema globularia, Intsia bijuga,
Memecylon edule and Pouteria linggensis typifying the flora of such a habitat.
Conservation status
Eleven species collected during this survey (inclusive of seagrasses) are listed in The
Red Book as ‘extinct’, 11 as endangered and 16 as vulnerable species (Table 3).
Blechnum orientale is relatively common on Chek Jawa, while the other
extinct, endangeredand vulnerable ferns are niche specific. The ‘extinct’ fern Adiantum
flabellulatum was found at one shady spot on a slope.
Of the mangrove species in the endangered and vulnerable categories,
Bruguiera parviflora is represented by a single tree, Sonneratia ovata by several
individuals, while Rhizophora stylosa is thriving and several saplings were seen around
the mother trees.
Litsea elliptica is relatively common. Archidendron ellipticum was found
growing in a sunny patch in the hill forest but saplings were not noted.
On the rocky shore, only one individual of Memecylon amplexicaule was
located. In contrast, Olea brachiata is common along forest margins on the seaward
side.
Aidia densiflora, a small tree, is common. A single specimen of Scaphium
macropodum was found. Garcinia hombroniana is represented by saplings along the
secondary forest pathways. Strobilanthes palawanensis, thought ‘extinct’, is ubiquitous
Survey of Chek Jawa, Singapore 281
Table 3. ‘Extinct’ , endangered and vulnerable species collected during the Chek Jawa
survey
‘Extinct’
Acanthaceae Strobilanthes palawanensis
Adiantaceae Adiantum flabellulatum
Cymodoceaceae Halodule pinifolia
Cymodoceaceae Halodule uninervis
Cymodoceaceae Thalassia hemprichii
Gnetaceae — Gnetum latifolium
Lauraceae Litsea myristicifolia
Myristicaceae Knema globularia
Palmae Calamus erinaceus
Pandanaceae Pandanus tetrodon
Rubiaceae Lasianthus hirsutus (as L. cyanocarpus)
Endangered
Guttiferae Garcinia hombroniana
Hydrocharitaceae Halophila beccarii
Hydrocharitaceae Halophila spinulosa
Leguminosae Archidendron ellipticum
Moraceae Artocarpus nitidus
Moraceae Ficus superba
Pteridaceae Pteris tripartita
Rhizophoraceae Bruguiera parviflora
Rubiaceae Diplospora malaccensis
Sonneratiaceae Sonneratia ovata
Schizaeaceae Schizaea dichotoma
Vulnerable
Blechnaceae Blechnum orientale
Celastraceae Salacia chinensis
Dracaenaceae Dracaena maingayi
Lauraceae Litsea elliptica
Lauraceae Litsea umbellata
Leguminosae Dalbergia pseudosissoo
Leguminosae Millettia pinnata
Loganiaceae Strychnos maingayi
Melastomataceae Memecylon amplexicaule
Meliaceae Xylocarpus moluccensis
Moraceae Ficus dubia
Oleaceae Olea brachiata
Rhizophoraceae Rhizophora stylosa
Rubiaceae Aidia densiflora
Schizaeaceae Lygodium circinnatum
Sterculiaceae Scaphium macropodum
282 Gard. Bull. Singapore 55 (2003)
on Chek Jawa. (It is possible that this species, which has pretty pink flowers, is a
garden escape). The seashore nutmeg, Knema globularia, is common on the seaward
side of coastal forest on Chek Jawa with several adult trees.
The first step in conservation would be to invest in a long-term monitoring
programme to follow the species flowering and fruiting seasons. Seeds should be
gathered and germinated so that saplings can be transplanted back into the surrounding
area. For example, Pandanus tetrodon was found on the steep slope in the coastal
forest is reperesented by a single huge plant with offshoots. Unless the offshoots are
transplanted elsewhere, it will remain the only plant on Chek Jawa. Even then the
gene pool for most of these rare plants must be rather depleted. Unless something is
done to increase the number of lone plants, the species will eventually become extinct.
Table 4. Specimens collected from Pulau Ubin before 1950 that occur on Chek
Jawa (Ex ‘Extinct’; V Vulnerable; R Rare; C Common)
Family Genus Species Status Collector No. Year collected
Adiantaceae Adiantum flabellutatum Ex Ridley, H.N. 5865 1892
Adiantaceae Pityrogramma calomelanos exotic Ridley, H.N. sn. 1894
Apocynaceae Willughbeia edulis R Ridley, H.N. 9501 1901
Asclepiadaceae Dischidia nummularia C Allen,BEGM. sn. 1949
Celastraceae Salacia chinensis V Ridley, H.N. sn. 1890
Compositae Pluchea indica R Hullett, R.W. 341 = 1884
Cyperaceae Fimbristylis ferruginea R Ridley, H.N. 5806 1892
Davalliaceae Davallia denticulata C Hullett, R.W. 50 1880
Dracaenaceae Dracaena maingayi V Hullett, R.W. sn. 1885
Euphorbiaceae Macaranga heynei C Furtado, C.X. 18338 1927
Euphorbiaceae Suregada multiflora R Ridley, H.N. 9497 1898
Leguminosae Derris trifoliata & Hullett, R.W. sn. 1885
Linaceae Indorouchera griffithiana R Allen,BE.G.M. — sn 1949
Loganiaceae Fagraea racemosa R Goodenough, JS. 1146 1890
Menispermaceae Fibraurea tinctoria R Ridley, H.N. sn 1893
Menispermaceae Tinospora crispa exotic Ridley, H.N. 1650 1890
Moraceae Artocarpus dadah R Ridley, H.N. 4721 1890
Moraceae Ficus kerkhovenii R Ridley, H.N. sn 1901
Moraceae Ficus variegata Se Ridley, H.N. sn. 1894
Ochnaceae Campylospermum | serratum R Ridley, H.N. sn 1897
Palmae Licuala spinosa R Ridley, H.N. sn 1894
Rhizophoraceae Bruguiera cylindrica R Ridley, H.N. 366 ~=61890
Rubiaceae Oxyceros longiflora R Ridley, H.N. 9487 1898
Rutaceae Clausena excavata R Ridley, H.N. 389 =. 1890
Sapindaceae Lepisanthes rubiginosa R Hullett, R.W. 386 =: 1884
Sapindaceae Mischocarpus sundaicus R Ridley, H.N. 9495 1890
Verbenaceae Clerodendrum inerme R Ridley, H.N. 373 = 1890
Verbenaceae Premna serratifolia R Ridley, H.N. sn 1893
Verbenaceae Vitex pinnata C Hullett, R.W. 373 ~=—- 1880
Survey of Chek Jawa, Singapore 283
Historic Collections
As mentioned above, it is not possible to determine if the specimens collected from
1800 to 1950 were collected from Chek Jawa as the locality was only recorded as
Pulau Ubin. Out of the 109 specimens from this period, only 29 species were collected
from Chek Jawa during this survey (Table 4) of which, Adiantum flabellulatum is
listed as ‘extinct’, and Salacia chinensis and Dracaena maingayi as vulnerable.
However, many other ‘extinct’, endangered and vulnerable species collected during
these early times were not seen on Chek Jawa (Table 5).
Conclusions
For its size, the flora of Chek Jawa is rich in biodiversity with 245 species in 171
genera and 80 families of vascular plants excluding alien species. This compares
with the survey for the whole of Pulau Ubin (Turner ef al., 1992), which listed 261
native species and 71 aliens. From the conservation point of view, with 11 species
listed as ‘extinct’, 11 as endangered and 16 as vulnerable, Chek Jawa is an important
conservation site and a significant component of Pulau Ubin flora. It has a good
representative flora of the seagrasses, the rocky shore and mangrove habitats. The
coastal hill, although heavily disturbed, is not secondary forest and harbours many of
the ‘extinct’, endangered or vulnerable species so conservation efforts should be
afforded to ensure its survival.
Acknowledgements
We thank Leni Duistermaat for her assistance in both collecting and identifying
Gramineae and Cyperaceae; to Ria Tan and her companions for their enthusiastic and
obliging help in the collection of seagrasses we missed; to Robert Teo and Choi Yook
Sau, Conservation Division at Pulau Ubin, for their assistance in facilitating the survey
as well as for the provision of logistics; to Saifuddin Suran, Biodiversity Centre,
NParks, for the map and data information pertaining to Chek Jawa. To Dr. I.M. Turner
for helpful amendments on the manuscript and for pointing out that Elephantopus
mollis is a new record and to Dr Ruth Kiew for guidance in conducting the field
surveys and in report writing.
284
Gard. Bull. Singapore 55 (2003)
Table 5. Vulnerable, endangered or ‘extinct’ species from old records that were not
found on Chek Jawa
‘Extinct’
Apocynaceae
Araliaceae
Commelinaceae
Connaraceae
Fagaceae
Leguminosae
Leguminosae
Loganiaceae
Orchidaceae
Orchidaceae
Rubiaceae
Endangered
Myrsinaceae
Myrsinaceae
Myrtaceae
Vulnerable
Annonaceae
Celastraceae
Dipterocarpaceae
Lauraceae
Melastomataceae
Palmae
Rhamnaceae
Rubiaceae
Selaginellaceae
Willughbeia flavescens
Schefflera lanceolata
Pollia secundiflora
Connarus planchonianus
Lithocarpus wallichianus
Aganope heptaphylla
Albizia retusa
Fagraea auriculata
Corymborkis veratrifolia
Thrixspermum calceolus
Coelospermum truncatum
Aegiceras corniculatum
Ardisia singaporensis
Tristaniopsis whiteana
Cyathostemma viridiflorum
Salacia viminea
Dipterocarpus sublamellatus
Actinodaphne macrophylla
Diplectria viminalis
Nenga pumila
Ventilago malaccensis
Ophiorrhiza singaporensis
Selaginella willdenowii
References
Chua, E.K. 2000. Pulau Ubin, Ours to Treasure. Simply Green, Singapore.
Chua, E.K. 2003. Chek Jawa, Discovering Singapore’s Biodiversity. Simply Green,
Singapore.
Survey of Chek Jawa, Singapore 285
Corlett, R.T. 1994. What is secondary forest? Journal of Tropical Ecology. 10: 445 —
447.
Corner, E.J.H. 1985. The botany of some islets east of Pahang and Johore. Gardens’
Bulletin Singapore. 38: 1-42
Ng, P.K.L. and Y.C. Wee 1994. The Singapore Red Data Book. Nature Society
(Singapore), Singapore.
Tan, R. and A. Yeo. 2003. Chek Jawa Guidebook. WildSingapore, Singapore.
The Straits Times. 2002. Chek Jawa: Anatomy of a U-turn. 2 January 2002.
The Straits Times. 2002. Reprieve for rustic Ubin. /5 January 2000.
The Straits Times. 2002. Plight of Ubin’s Kampung folk. 24 August 2002.
Turner, I.M., H.T.W. Tan, K.S Chua, A. Samsuri and Y.C. Wee. 1992. A botanical
survey of Pulau Ubin. Gardens’ Bulletin Singapore. 44: 51-71
URA Development Guide Plans. 2003. http://www.ura.gov.sg/landuse/devtguide-
frame.htm (Accessed 24 October 2003).
Appendix
Checklist of Vascular Plants Collected from Chek Jawa
[Ex] Extinct; [En] Endangered; [V] Vulnerable and [R] Rare; [exotic] plants not
originating in the region and [C] Common status from The Singapore Red Data Book
(1994).
Ferns
Adiantaceae
Adiantum flabellulatum L.; habit : herb; occurrence: rare: habitat : coastal forest;
conservation status: Ex; specimen(s): Gwee, A.T. GAT 92.
Adiantum latifolium Lam.; habit : herb; 0.75m tall; occurrence: frequent; habitat:
coastal forest; conservation status: exotic; specimen(s): Gwee, A.T. GAT 145.
Pityrogramma calomelanos (L.) Link; habit : herb; 0.5m tall; habitat: wasteland;
conservation status: exotic; specimen(s): Gwee, A.T. GAT 148.
286 Gard. Bull. Singapore 55 (2003)
Taenitis blechnoides (Willd.) Sw.; habit : herb; 0.3m tall; occurrence: locally frequent;
habitat: coastal forest; conservation status: common; specimen(s): Gwee, A.T. GAT
187.
Blechnaceae
Blechnum orientale L.: habit : herb: 1.5m tall; habitat : coastal forest: conservation
status: V; specimen(s): Gwee, A.T. GAT 167.
Stenochlaena palustris (Burm.f.) Bedd.; habit : herb; occurrence: frequent; habitat :
coastal forest; conservation status: common; specimen(s): Gwee, A.T. GAT 210.
Davalliaceae
Davallia denticulata (Burm .f.) Mett.ex Kuhn; habit : epiphytic herb; habitat: coastal
forest; conservation status: common; specimen(s): Gwee, A.T. GAT 57.
Dennstaedtiaceae
Pteridium esculentum (G. Forst.) Cockayne; habit : herb; habitat: sandy shore;
specimen(s): Gwee, A.T. GAT 316.
Oleandraceae
Nephrolepis auriculata (L.) Trimen; habit : herb; occurrence: frequent; habitat :
wasteland / coastal hill; conservation status: common; specimen(s): Gwee, A.T. GAT
Ha B
Polypodiaceae :
Drynaria quercifolia (L.) J.Sm.; habit : epiphytic herb; habitat : coastal forest;
conservation status: common; specimen(s): Gwee, A.T. GAT 44.
Microsorum scolopendrium (Burm.f.) Copel.; habit : epiphytic herb; habitat: coastal
forest; specimen(s): Gwee, A.T. GAT 173.
Pyrrosia nummulariifolia (Sw.) Ching; habit : climber; habitat : coastal forest;
specimen(s): Gwee, A.T. GAT 351.
Pyrrosia piloselloides (L.) M.G. Price; habit : climber; habitat: coastal forest;
conservation status: common; specimen(s): Gwee, A.T. GAT 199.
Pteridaceae
Acrostichum aureum L.; 2.5 tall; habitat: mangrove; conservation status: common:
specimen(s): Gwee, A.T. GAT 152.
Survey of Chek Jawa, Singapore 287
Acrostichum speciosum Willd.; habit : herb; habitat: mangrove; conservation status:
common; specimen(s): Gwee, A.T. GAT 283.
Pteris ensiformis Burm.f.; habit : herb; habitat: coastal forest; conservation status:
common; specimen(s): Leong, P. PL 22.
Pteris tripartita Sw.; habit: herb; habitat: coastal forest; conservation status: En;
specimen(s): Gwee, A.T. GAT 186.
Schizaeaceae
Lygodium circinnatum (Burm.f.) Sw.; habit : herb; habitat: wasteland; conservation
status: V; specimen(s): Leong, P. PL 8.
Lygodium salicifolium C. Presl; habit: climber; habitat : coastal forest; conservation
status: common; specimen(s): Gwee, A.T. GAT 335.
Schizaea dichotoma (L.) J. Sm.; habit : herb; habitat : coastal forest; conservation
status: En; specimen(s): Gwee, A.T. GAT 340.
Schizaea digitata (L.) Sw.; habit : herb; specimen(s): Gwee, A.T. GAT 292.
Gymnosperms
Gnetaceae
Gnetum gnemon L.; habit : tree; habitat : wasteland; conservation status: R;
specimen(s): Leong, P. PL 3.
Gnetum latifolium Blume; habit : climber; habitat: coastal forest; conservation status:
Ex; specimen(s): Gwee, A.T. GAT 222.
Gnetum microcarpum Blume; habit : climber; habitat: coastal forest; conservation
status: R; specimen(s): Gwee, A.T. GAT 64.
Monocotyledons
Colchicaceae
Gloriosa superba L.; habit: climber; habitat: wasteland: conservation status: exotic:
specimen(s): Gwee, A.T. GAT 252.
Cymodoceaceae
Cymodocea rotundata Ehrenb. & Hempr.; habit : seagrass; habitat: intertidal zone;
conservation Status: R; specimen(s): Gwee, A.T. GAT 326.
288 Gard. Bull. Singapore 55 (2003)
Halodule pinifolia (Miki) Hartog; habit: seagrass; habitat: intertidal zone;
conservation status: Ex; specimen(s): Gwee, A.T. GAT 325.
Halodule uninervis (Forssk.) Asch.; habit : seagrass; habitat: intertidal zone;
conservation status: Ex; specimen(s): Tan, R. et al. GAT 398.
Cyperaceae
Cyperus cyperoides (L.) Kuntze; habit : herb; occurrence: frequent; habitat: wasteland;
conservation status: R; specimen(s): Leong, P. PL 23.
Cyperus imbricatus Retz.; habit : herb; 1m tall; habitat : wasteland; conservation
status: R; specimen(s): Gwee, A.T. GAT 154.
Cyperus Javanicus Houtt.; habit : herb; habitat: wasteland; conservation status: exotic;
specimen(s): Gwee, A.T. GAT 264.
Cyperus trialatus (Boeck.) Kern; habit : herb; 0.3m tall; habitat : wasteland;
conservation status: R; specimen(s): Gwee, A.T. GAT 190.
Fimbristylis ferruginea (L.) Vahl; habit : herb; occurrence: locally frequent; habitat:
wasteland; conservation status: R; specimen(s): Gwee, A.T. GAT 263.
Kyllinga polyphylla Willd. ex Kunth; habit : herb; habitat : wasteland; specimen(s):
Gwee,A.T..GAT 150.
Mapania enodis (Miq.) C.B. Clarke; habit: herb; occurrence: locally frequent; habitat:
coastal forest; conservation status: R; specimen(s):;Gwee, A.T. GAT 96.
Scleria ciliaris Nees; habit: herb; habitat: wasteland; conservation status: common;
specimen(s): Gwee, A.T. GAT 297.
Dracaenaceae
Dracaena angustifolia Roxb.; habit : shrub; habitat: coastal forest; specimen(s): Gwee,
A.T. GAT 253.
Dracaena maingayi Hook.f.; habit : tree; habitat: coastal forest; conservation status:
V; specimen(s): Gwee, A.T. GAT 55.
Flagellariaceae
Flagellaria indica L.; habit: climber; occurrence: locally frequent; habitat: mangrove:
conservation status: R; specimen(s): Gwee, A.T. GAT 192.
Survey of Chek Jawa, Singapore 289
Gramineae
Bambusa vulgaris Schrad.; habitat: coastal forest; conservation status: R; specimen(s):
Gwee, A.T. GAT 334.
Centotheca lappacea (L.) Desv.; habit : herb; 0.35m tall; habitat : wasteland;
conservation status: R; specimen(s): Gwee, A.T. GAT 159.
Mnesithea glandulosa (Trin.) de Koning & Sosef; habit : herb; 1.5m tall; habitat :
wasteland; conservation status: R; specimen(s): Gwee, A.T. GAT 147.
Panicum nodosum Kunth; habit : herb; habitat : wasteland; specimen(s): Gwee, A.T.
GAT 12.
Thysanolaena latifolia (Roxb. ex Hornem.) Honda; habit : herb; habitat: wasteland;
specimen(s): Gwee, A.T. GAT 345.
Hydrocharitaceae
Halophila beccarii Asch.; habit : seagrass; habitat: intertidal zone; conservation status:
En; specimen(s): Tan, R. et al. GAT 396.
Halophila ovalis (R.Br.) Hook.f.; habit : seagrass; occurrence: frequent; habitat :
intertidal zone; conservation status: R; specimen(s): Gwee, A.T. GAT 324.
Halophila spinulosa (R.Br.) Asch.; habit : seagrass; occurrence: frequent; habitat :
intertidal zone; conservation status: En; specimen(s): Gwee, A.T. GAT 322.
Thalassia hemprichii (Ehrenb.) Asch.; habit : seagrass; habitat: intertidal zone;
conservation status: Ex; specimen(s): Tan, R. et al. GAT 393.
Hypoxidaceae
Molineria latifolia (Dryand.) Herb. ex Kurz var. latifolia; habit: herb; habitat: coastal
forest; conservation status: R; specimen(s): Gwee, A.T. GAT 68.
Orchidaceae
Dendrobium crumenatum Sw.; habit : herb; habitat: rocky shore; conservation status:
common; specimen(s): Gwee, A.T. GAT 171.
Eulophia graminea Lindl.; habit : herb; habitat : sandy shore; conservation status:
common; specimen(s): Gwee, A.T. GAT 268.
290 Gard. Bull. Singapore 55 (2003)
Palmae
Calamus erinaceus (Becc.) J. Dransf.; habit : rattan; habitat: coastal forest;
conservation status: Ex; specimen(s): Gwee, A.T. GAT 53.
Daemonorops sepal Becc.; habit : rattan; habitat: coastal forest; specimen(s): Gwee,
A.T. GAT 343.
Licuala spinosa Wurmb; habit : shrub; 3m tall; habitat: coastal forest; conservation
status: R; specimen(s): Gwee, A.T. GAT 9.
Nypa fruticans Wurmb; habitat: mangrove; conservation status: R; specimen(s):
Gwee, A.T. GAT 317.
Oncosperma tigillarium (Jack) Ridl.; habit : tree; habitat : coastal forest; conservation
status: R; specimen(s): Gwee, A.T. GAT 385.
Pandanaceae
Pandanus odoratissimus L.£.; habitat : mangrove; conservation status: common;
specimen(s): Gwee, A.T. GAT 266.
Pandanus tetrodon Rid1.; habitat : coastal forest; conservation status: Ex; specimen(s):
Gwee, A.T. GAT 392.
Phormiaceae
Dianella ensifolia (L.) DC.; habit : herb; occurrence: frequent; habitat: coastal forest;
conservation status: common; specimen(s): Gwee, A.T. GAT 332.
Smilacaceae
Smilax megacarpa A. DC. & C. DC.; habit : climber; habitat: coastal forest;
conservation status: R; specimen(s): Gwee, A.T. GAT 79.
Smilax setosa Miq.; habit : climber; habitat: coastal forest; conservation status:
common; specimen(s): Gwee, A.T. GAT 75.
Zingiberaceae
Zingiber griffithii Baker; habit : herb; 0.3m tall; habitat: coastal forest; conservation
status: R; specimen(s): Gwee, A.T. GAT 160.
Dicotyledons
Acanthaceae
Survey of Chek Jawa, Singapore 291
Strobilanthes palawanensis Elmer; habit : herb; 0.75m tall; occurrence: locally
frequent; habitat : wasteland; conservation status: Ex; specimen(s): Gwee, A.T. GAT
146.
Aizoaceae
Sesuvium portulacastrum (L.) L.; habit: herb; habitat: wasteland; conservation status:
R; specimen(s): Gwee, A.T. GAT 225.
Alangiaceae
Alangium javanicum (Blume) Wangerin; habit: tree; 3m tall; habitat: wasteland;
specimen(s): Gwee, A.T. GAT 110.
Amaranthaceae
Cyathula prostrata (L.) Blume; habit : herb; habitat: wasteland; conservation status:
R; specimen(s): Gwee, A.T. GAT 347.
Anacardiaceae
Buchanania sessifolia Blume; habit : tree; 1.5m tall; habitat: coastal forest;
conservation status: R; specimen(s): Gwee, A.T. GAT 184.
Mangifera indica L.; habit : tree; 20m tall; habitat: coastal forest; conservation status:
exotic; specimen(s): Gwee, A.T. GAT 76.
Annonaceae
Artabotrys suaveolens (Blume) Blume; habit : climber; occurrence: frequent; habitat: »
coastal forest; conservation status: R; specimen(s): Gwee, A.T. GAT 183.
Desmos chinensis Lour.; habit : tree; occurrence: frequent; habitat: coastal forest;
conservation status: R; specimen(s): Gwee, A.T. GAT 63.
Melodorum aberrans (Maingay ex Hook.f. & Thomson) J. Sm.; habit : climber;
habitat: coastal forest; specimen(s): Gwee, A.T. GAT 196.
Phaeanthus ophthalmicus (Roxb. ex G. Don) J. Sinclair; habit: tree: habitat : coastal
forest; conservation status: R; specimen(s): Gwee, A.T. GAT 276.
Popowia fusca King; habit : tree; 0.75m tall; habitat: coastal forest; conservation
status: R; specimen(s): Gwee, A.T. GAT 130.
Uvaria cordata (Dunal) Alston; habit: climber; 1m tall; habitat: coastal forest;
conservation Status: R; specimen(s): Gwee, A.T. GAT 124.
292 Gard. Bull. Singapore 55 (2003)
Uvaria grandiflora Roxb. ex Hornem. var. grandiflora; habitat : coastal forest;
conservation status: R; specimen(s): Gwee, A.T. GAT 386.
Uvaria hirsuta Jack; habit: climber; habitat: coastal forest; conservation status: R:
specimen(s): Gwee, A.T. GAT 337.
Xylopia malayana Hook.f. & Thomson; habit : tree; habitat : coastal forest;
conservation status: R; specimen(s): Gwee, A.T. GAT 302.
Apocynaceae
Cerbera manghas L.; habit: tree; habitat : mangrove; conservation status: R:
specimen(s): Gwee, A.T. GAT 281.
Willughbeia edulis Roxb.; habit : climber; occurrence: locally frequent; habitat :
coastal forest; conservation status: R; specimen(s): Gwee, A.T. GAT 106.
Aquifoliaceae
Ilex cymosa Blume; habit : tree; 5m tall; habitat : wasteland; conservation status: R:
specimen(s): Gwee, A.T. GAT 206.
Asclepiadaceae
Dischidia major (Vahl) Merr.; habit : epiphytic herb; occurrence: frequent; habitat:
coastal forest; conservation status: R; specimen(s): Leong, P. PL 16.
Dischidia nummularia R.Br.; habit : epiphytic herb; occurrence: frequent; habitat:
coastal forest; conservation status: common; specimen(s): Gwee, A.T. GAT 10.
Hoya verticillata (Vahl) G. Don var. verticillata; habit: climber; occurrence: frequent;
habitat : coastal forest; conservation status: common; specimen(s): Gwee, A.T. GAT
109.
Avicenniaceae
Avicennia alba Blume; habit : tree; 6m tall; occurrence: locally frequent; habitat:
mangrove; conservation status: common; specimen(s): Gwee, A.T. GAT 117.
Avicennia officinalis L.; habit : tree; 4m tall; occurrence: frequent; habitat : mangrove;
conservation status: R; specimen(s): Gwee, A.T. GAT 226.
Avicennia rumphiana Hallier f.; habit : tree; 4m tall; habitat : mangrove; conservation
status: R; specimen(s): Gwee, A.T. GAT 235.
Survey of Chek Jawa, Singapore 293
Boraginaceae
Cordia curassavica (Jacq.) Roem. & Schult.; habit : shrub; 1m tall; habitat: sandy
shore; conservation status: common; specimen(s): Gwee, A.T. GAT 174.
Burseraceae
Dacryodes rostrata (Blume) H.J. Lam; habit : tree; 25m tall; habitat : coastal forest;
conservation status: R; specimen(s): Gwee, A.T. GAT 158.
Celastraceae
Salacia chinensis L.; habit : climber; habitat: mangrove; conservation status: V;
specimen(s): Gwee, A.T. GAT 86.
Salacia korthalsiana Migq.; habit : climber; habitat: coastal forest; conservation status:
R; specimen(s): Gwee, A.T. GAT 387.
Chrysobalanaceae
Licania splendens (Korth.) Prance; habit : tree; habitat: coastal forest; conservation
status: R; specimen(s): Gwee, A.T. GAT 309.
Maranthes corymbosa Blume; habit : tree; occurrence: locally frequent; habitat :
coastal forest; conservation status: R; specimen(s): Gwee, A.T. GAT 320.
Combretaceae
Combretum sundaicum Mig.; habit : climber; habitat: coastal forest; conservation
status: R; specimen(s): Gwee, A.T. GAT 376.
Lumnitzera littorea (Jack) Voigt; habit : tree; habitat: mangrove; conservation Status:
R; specimen(s): Gwee, A.T. GAT 287.
Terminalia catappa L.; habit : tree; habitat: sandy shore; conservation status: common;
specimen(s): Gwee, A.T. GAT 260.
Compositae
Ageratum conyzoides L.; habit : herb; occurrence: frequent; habitat: wasteland;
conservation status: exotic; specimen(s): Gwee, A.T. GAT 348.
Elephantopus mollis Kunth; habit: herb; 0.75m tall; occurrence: frequent; habitat :
wasteland; specimen(s): Gwee, A.T. GAT 195.
294 Gard. Bull. Singapore 55 (2003)
Pluchea indica (L.) Less.; habit : shrub; occurrence: frequent; habitat : wasteland;
conservation status: R; specimen(s): Gwee, A.T. GAT 229.
Synedrella nodiflora (L.) Gaertn.; habit: herb; occurrence: frequent; habitat:
wasteland; conservation status: exotic; specimen(s): Gwee, A.T. GAT 363.
Wollastonia biflora (L.) DC.; habit : herb; specimen(s): Leong, P. PL 13.
Connaraceae
Cnestis palala (Lour.) Merr.; habit : climber; occurrence: locally frequent; habitat:
coastal forest; conservation status: R; specimen(s): Gwee, A.T. GAT 126.
Rourea fulgens Planch.; habit : climber; habitat: coastal forest; conservation status:
R; specimen(s): Gwee, A.T. GAT 99.
Rourea mimosoides (Vahl) Planch.; habit: climber; habitat : coastal forest:
conservation status: R; specimen(s): Gwee, A.T. GAT 122.
Rourea minor (Gaertn.) Leenh.; habit : climber; habitat: coastal forest; conservation
status: R; specimen(s): Gwee, A.T. GAT 375.
Convolvulaceae
Erycibe malaccensis C.B. Clarke; habit : climber; occurrence: locally frequent; habitat:
coastal forest; conservation status: R; specimen(s): Gwee, A.T. GAT 277.
Erycibe tomentosa Blume; habit : climber; occurrence: locally frequent; habitat :
coastal forest; conservation status: common; specimen(s): Gwee, A.T. GAT 54.
Cucurbitaceae
Coccinea grandis (L.) Voigt; habit : climber; habitat: wasteland; conservation status:
exotic; specimen(s): Leong, P. PL 11.
Dilleniaceae
Tetracera akara (Burm.f.) Merr.; habit : climber; habitat : coastal forest; specimen(s):
Gwee, A.T. GAT 127.
Tetracera indica (Christm. & Panz.) Merr.; habit : climber; habitat: coastal forest;
conservation status: common; specimen(s): Gwee, A.T. GAT 4.
Tetracera macrophylla Wall. ex Hook.f. & Thomson; habit : climber; habitat : coastal
forest; conservation status: R; specimen(s): Gwee, A.T. GAT 306.
Survey of Chek Jawa, Singapore 295
Ebenaceae
Diospyros ferrea (Willd.) Bakh.; habit : tree; 5m tall; occurrence: locally frequent;
habitat: rocky shore; conservation status: R; specimen(s): Gwee, A.T. GAT 177.
Diospyros lanceifolia Roxb.; habit : tree; habitat : coastal forest; conservation status:
R; specimen(s): Leong, P. PL 18.
Elaeocarpaceae
Elaeocarpus pedunculatus Wall. ex Mast.; habit : tree; 2.5m tall; habitat: coastal
forest; conservation status: R; specimen(s): Gwee, A.T. GAT 157.
Erythroxylaceae
Erythroxylum cuneatum (Miq.) Kurz; habit: tree; habitat : coastal forest; conservation
status: R; specimen(s): Gwee, A.T. GAT 344.
Euphorbiaceae
Aporusa frutescens Blume; habit : tree; habitat: coastal forest; conservation status:
R; specimen(s): Gwee, A.T. GAT 298.
Breynia racemosa (Blume) Miill.Arg.; habit : shrub; habitat : coastal forest;
conservation status: common; specimen(s): Gwee, A.T. GAT 205.
Croton hirtus L Hér.; habit : shrub; habitat : coastal forest; conservation status: exotic;
specimen(s): Gwee, A.T. GAT 114.
Excoecaria agallocha L.; habit: tree; habitat: mangrove; conservation status: R;
specimen(s): Gwee, A.T. GAT 224.
Glochidion littorale Blume; habit : tree; habitat: coastal forest; conservation status:
R; specimen(s): Gwee, A.T. GAT 296.
Glochidion superbum Baill.; habit : tree; 5m tall; occurrence: locally frequent; habitat:
coastal forest; conservation status: R; specimen(s): Gwee, A.T. GAT 131.
Macaranga heynei 1.M. Johnst.; habit : tree; 3m tall; habitat: coastal forest;
conservation status: common; specimen(s): Gwee, A.T. GAT 5.
Macaranga hypoleuca (Rchb.f. & Zoll.) Miill.Arg.; habit : tree; 3m tall; habitat:
coastal forest; conservation status: common; specimen(s): Gwee, A.T. GAT 189.
296 Gard. Bull. Singapore 55 (2003)
Macaranga lowii King ex Hook.f.; habit : tree; 3m tall; habitat: coastal forest;
conservation status: R; specimen(s): Gwee, A.T. GAT 77.
Triadica cochinchinensis Lour.; habit : tree; 4m tall; habitat: coastal forest:
conservation status: common; specimen(s): Gwee, A.T. GAT 133.
Sauropus androgynus (L.) Merr.; habit : shrub; habitat : coastal forest; conservation
status: common; specimen(s): Gwee, A.T. GAT 6.
Suregada multiflora (Juss.) Baill.; habit : tree; occurrence: locally frequent; habitat:
coastal forest; conservation status: R; specimen(s): Gwee, A.T. GAT 164.
Flacourtiaceae
Flacourtia inermis Roxb.; habit : tree; habitat : wasteland; specimen(s): Gwee, A.T.
GAT 207.
Goodeniaceae
Scaevola taccada (Gaertn.) Roxb.; habit : shrub; habitat: rocky shore; conservation
status: common; specimen(s): Gwee, A.T. GAT 358.
Guttiferae
Garcinia atroviridis Griff. ex T. Anderson; habit : tree; habitat: coastal forest:
conservation status: R; specimen(s): Gwee, A.T. GAT 275.
Garcinia hombroniana Pierre; habit : tree; 8m tall; habitat: rocky shore; conservation
status: En; specimen(s): Gwee, A.T. GAT 141.
Garcinia parvifolia (Miq.) Migq.; habit : tree; habitat : coastal forest; conservation
status: R; specimen(s): Gwee, A.T. GAT 274.
Icacinaceae
Gonocaryum gracile Miq.; habit : tree; 6m tall; habitat : coastal forest; conservation
status: R; specimen(s): Gwee, A.T. GAT 129.
Ixonanthaceae
Ixonanthes icosandra Jack; habit : tree; habitat: coastal forest; conservation status:
R; specimen(s): Gwee, A.T. GAT 291.
Labiatae
Hyptis brevipes Poit.; habit : herb; occurrence: frequent; habitat : wasteland;
conservation status: exotic; specimen(s): Leong, P. PL 12.
Survey of Chek Jawa, Singapore 297
Lauraceae
Cinnamomum iners Reinw.: habit : tree: 6m tall: habitat: coastal forest: conservation
status: common; specimen(s): Gwee, A.T. GAT 70.
Litsea elliptica Blume; habit : tree; 5m tall; habitat : coastal forest; conservation
status: V; specimen(s): Gwee, A.T. GAT 188.
Litsea myristicifolia (Wall. ex Nees) Hook.f.; habit : tree; 1m tall; habitat: coastal
forest; conservation status: Ex; specimen(s): Gwee, A.T. GAT 185.
Litsea umbellata (Lour.) Merr.; habit: tree; 2m tall; habitat: coastal forest; conservation
status: V; specimen(s): Gwee, A.T. GAT 93.
Neolitsea zeylanica (Nees) Merr.; habit : tree; 15m tall; occurrence: frequent; habitat:
coastal forest; conservation status: R; specimen(s): Leong, P. PL 2.
Leguminosae
Adenanthera malayana Kosterm.; habit : tree; habitat: coastal forest; conservation
status: R; specimen(s): Gwee, A.T. GAT 381.
Aganope thyrsiflora (Benth.) Polhill; habit: climber; habitat : rocky shore;
conservation status: R; specimen(s): Gwee, A.T. GAT 82.
Archidendron contortum (Martelli) 1.C. Nielsen; habit : tree; occurrence: locally
frequent; habitat: coastal forest; conservation status: R; specimen(s): Gwee, A.T. GAT
119.
Archidendron ellipticum (B1.) Nielsen; habit : tree; habitat: coastal forest;
conservation status: V; specimen(s): Gwee, A.T. GAT 308.
Archidendron jiringa (Jack) 1.C. Nielsen; habit : tree: habitat: coastal forest;
conservation status: R; specimen(s): Gwee, A.T. GAT 198.
Canavalia cathartica Thouars; habit : climber: habitat : wasteland: conservation
status: common; specimen(s): Gwee, A.T. GAT 212.
Canavalia rosea (Sw.) DC.: habit : climber; habitat: sandy shore; conservation status:
common; specimen(s): Gwee, A.T. GAT 250.
Clitoria pubescens; habit: climber; habitat: sandy shore: specimen(s): Leong, P. PL
yA
298 Gard. Bull. Singapore 55 (2003)
Dalbergia candenatensis (Dennst.) Prain; habit : climber; occurrence: frequent;
habitat: coastal forest; conservation status: R; specimen(s): Gwee, A.T. GAT 179.
Dalbergia pseudosissoo Miq.; habit : climber; habitat : coastal forest; conservation
status: V; specimen(s): Gwee, A.T. GAT 120.
Dalbergia velutina Benth.; habit : climber; habitat: coastal forest; conservation status:
R; specimen(s): Gwee, A.T. GAT 216.
Dendrolobium umbellatum (L.) Benth.; habit : tree; 1.5m tall; habitat: wasteland;
conservation status: common; specimen(s): Gwee, A.T. GAT 115.
Derris amoena Benth. var. amoena; habit : climber; occurrence: frequent; habitat :
coastal forest; specimen(s): Gwee, A.T. GAT 355.
Derris trifoliata Lour.; habit : tree; habit : climber; habitat: coastal forest; conservation
status: common; specimen(s): Gwee, A.T. GAT 162.
Desmodium heterocarpon (L.) DC. ssp. heterocarpon; habit : shrub; habitat:
wasteland; conservation status: R; specimen(s): Gwee, A.T. GAT 373.
Entada spiralis Ridl\.; habit: climber; occurrence: frequent; habitat : coastal forest;
conservation status: R; specimen(s): Gwee, A.T. GAT 8.
Intsia bijuga (Colebr.) Kuntze; habit : tree; 9m tall; occurrence: rare; habitat : rocky
shore; conservation status: R; specimen(s): Gwee, A.T. GAT 138.
Millettia pinnata (L.) G. Panigrahi; habit : tree; habitat: sandy shore; conservation
status: V; specimen(s): Gwee, A.T. GAT 241.
Linaceae
Indorouchera griffithiana (Planch.) Hallier f.; habit : climber; habitat: coastal forest;
conservation status: R; specimen(s): Gwee, A.T. GAT 1.
Loganiaceae
Fagraea fragrans Roxb.; habit: tree; habitat : coastal forest; conservation status:
common; specimen(s): Gwee, A.T. GAT 61.
Fagraea racemosa Jack ex Wall.; habit : tree; 4m tall; occurrence: frequent; habitat:
coastal forest; conservation status: R; specimen(s): Gwee, A.T. GAT 217.
Survey of Chek Jawa, Singapore 299
Strychnos ignatii Berg.; habit: climber; occurrence: frequent; habitat : coastal forest;
conservation status: R; specimen(s): Gwee, A.T. GAT 95.
Strychnos maingayi C.B. Clarke; habit : climber; habitat: coastal forest; conservation
status: V; specimen(s): Gwee, A.T. GAT 285.
Malvaceae
Sida acuta Burm. f.; habit: shrub; habitat: wasteland; conservation status: common;
specimen(s): Gwee, A.T. GAT 153.
Sida rhombifolia L.; habit: shrub; 1m tall; habitat: wasteland; conservation status:
common; specimen(s): Gwee, A.T. GAT 111.
Urena lobata L.; habit: shrub; occurrence: frequent; habitat: wasteland; conservation
status: common; specimen(s): Gwee, A.T. GAT 156.
Melastomataceae
Clidemia hirta (L.) D. Don; habit : shrub; occurrence: frequent; habitat: wasteland;
conservation status: exotic; specimen(s): Gwee, A.T. GAT 67.
Memecylon amplexicaule Roxb.; habit : tree; habitat: coastal forest; conservation
status: V; specimen(s): Gwee, A.T. GAT 338.
Memecylon edule Roxb. var. edule; habit : tree; 6m tall; habitat: rocky shore;
conservation status: R; specimen(s): Gwee, A.T. GAT 65.
Memecylon edule Roxb. var. ovatum (Sm.) C.B. Clarke; habit : tree; 7m tall; habitat:
rocky shore; conservation status: R; specimen(s): Gwee, A.T. GAT 74.
Meliaceae
Xylocarpus granatum J. Konig; habit : tree; 10m tall; habitat: mangrove; conservation
status: R; specimen(s): Gwee, A.T. GAT 140.
Xylocarpus moluccensis (Lam.) M. Roem.; habit : tree; habitat: mangrove;
conservation status: V; specimen(s): Gwee, A.T. GAT 254.
Menispermaceae
Fibraurea tinctoria Lour.; habit : climber; occurrence: frequent; habitat: coastal forest;
conservation Status: R; specimen(s): Gwee, A.T. GAT 56.
300 Gard. Bull. Singapore 55 (2003)
Tinospora crispa (L.) Hook.f. & Thomson; habit : climber; habitat: wasteland;
conservation status: exotic; specimen(s): Gwee, A.T. GAT 112.
Moraceae
Artocarpus dadah Miq.; habit : tree; 9-10m seen tall; habitat : coastal forest;
conservation status: R; specimen(s): Gwee, A.T. GAT 219.
Artocarpus nitidus Trécul ssp. griffithiti (King) F.M. Jarrett; habit : tree; habitat:
coastal forest; conservation status: En; specimen(s): Gwee, A.T. GAT 295.
Ficus aurantiacea Griff.; habit : climber; habitat: coastal forest; conservation status:
R; specimen(s): Gwee, A.T. GAT 204.
Ficus dubia Wall. ex King; habit : tree; habitat: coastal forest; conservation status:
V; specimen(s): Gwee, A.T. GAT 104.
Ficus grossularioides Burm. £.; habit: tree; habitat : coastal forest; conservation status:
common; specimen(s): Gwee, A.T. GAT 78.
Ficus heteropleura Blume; habit : tree; habitat : coastal forest; conservation status:
common; specimen(s): Gwee, A.T. GAT 203.
Ficus kerkhovenii Valeton; habit : tree; habitat: coastal forest; conservation status:
R; specimen(s): Gwee, A.T. GAT 270.
Ficus microcarpa L..; habit : tree; 1m tall; habitat: sandy shore; conservation status:
common; specimen(s): Gwee, A.T. GAT 169.
Ficus superba (Miq.) Miq.; habit : tree; habitat: sandy shore; conservation status:
En; specimen(s): Gwee, A.T. GAT 259.
Ficus variegata Blume; habit : tree; 7m tall; habitat : coastal forest; conservation
status: common; specimen(s): Gwee, A.T. GAT 113.
Ficus vasculosa Wall. ex Miq.; habit : tree; habitat: coastal forest; conservation Status:
R; specimen(s): Gwee, A.T. GAT 52.
Ficus villosa Blume; habit : climber; habitat : coastal forest; conservation status: R;
specimen(s): Gwee, A.T. GAT 197.
Survey of Chek Jawa, Singapore 301
Myristicaceae
Knema globularia (Lam.) Warb.; habit : tree; 4-18m tall; habitat: rocky shore/ coastal
hill; conservation status: Ex; specimen(s): Gwee, A.T. GAT 102.
Myrsinaceae
Ardisia crenata Sims; habit : shrub; habitat : coastal forest; conservation status: R;
specimen(s): Gwee, A.T. GAT 202.
Ardisia crispa A.DC.; habit : tree; 1m tall; habitat: coastal forest; specimen(s): Gwee,
A.T. GAT 2.
Embelia ribes Burm.; habit: climber; occurrence: frequent; habitat : coastal forest;
conservation Status: R; specimen(s): Leong, P. PL 5.
Myrtaceae
Psidium cattleianum, habit: tree; habitat : wasteland; specimen(s): Gwee, A.T. GAT
STZ.
Rhodamnia cinerea Jack; habit : tree; habitat : coastal forest; conservation status:
common; specimen(s): Gwee, A. T. GAT 69.
Syzygium lineatum (DC.) Merr. & L.M. Perry; habit : tree; occurrence: frequent;
habitat: coastal forest; conservation status: common; specimen(s): Gwee, A.T. GAT
89.
Syzygium zeylanicum (L.) DC.; habit : tree; 3.5m tall; occurrence: frequent; habitat:
coastal forest; conservation status: common; specimen(s): Gwee, A.T. GAT 249.
Ochnaceae
Campylospermum serratum (Gaertn.) Bittrich & M.C.E. Amaral; habit : tree;
occurrence: frequent; habitat: coastal forest; conservation status: R; specimen(s):
Gwee,A.T. GAT 81.
Olacaceae
Ximenia americana L.; habit : shrub; 4m tall; habitat: mangrove; conservation status:
R; specimen(s): Gwee, A.T. GAT 245.
Oleaceae
Jasminum elongatum (Bergius) Willd.; habit : climber; habitat : coastal forest;
conservation status: exotic; specimen(s): Leong, P. PL 7.
302 Gard. Bull. Singapore 55 (2003)
Olea brachiata (Lour.) Merr.; habit : tree; 2m tall; occurrence: frequent; habitat :
coastal forest; conservation status: V; specimen(s): Gwee, A.T. GAT 72.
Onagraceae
Ludwigia hyssopifolia (G. Don) Exell; habit : herb; 0.5m tall; habitat: wasteland;
conservation status: common; specimen(s): Gwee, A.T. GAT 194.
Opiliaceae
Champereia manillana (Blume) Merr.; habit : tree; 4m tall; occurrence: frequent;
habitat : coastal forest; conservation status: common; specimen(s): Gwee, A.T. GAT
118.
Pandaceae
Microdesmis caseariifolia Planch.; habit : tree; 4m tall; occurrence: frequent; habitat:
coastal forest; conservation status: R; specimen(s): Gwee, A.T. GAT 128.
Passifloraceae
Adenia macrophylla (Blume) Koord. var. macrophylla; habit : climber; habitat:
wasteland; conservation status: R; specimen(s): Gwee, A.T. GAT 368.
Passiflora foetida L.; habit : climber; habitat : wasteland; conservation status: exotic;
specimen(s): Gwee, A.T. GAT 193.
Passiflora laurifolia L.; habit : climber; occurrence: frequent; habitat: wasteland;
conservation status: exotic; specimen(s): Gwee, A.T. GAT 59.
Polygalaceae
Polygala paniculata L.; habit : herb; habitat: wasteland; conservation status: exotic;
specimen(s): Gwee, A.T. GAT 367.
Polygonaceae
Muelenbeckia platyclados; habit: herb; habitat : wasteland; specimen(s): Gwee,A.T.
GAT 228.
Portulacaceae
Portulaca pilosa L.; habit : herb; 0.1m tall; habitat : rocky shore; conservation status:
exotic; specimen(s): Leong, P. PL 10.
Rhamnaceae
Colubrina asiatica L. ex Brongn.; habit : shrub; habitat: sandy shore; conservation
status: R; specimen(s): Gwee, A.T. GAT 243.
Survey of Chek Jawa, Singapore 303
Rhizophoraceae
Bruguiera cylindrica (L.) Blume; habit : tree; 2.5m tall; occurrence: frequent; habitat:
mangrove; conservation status: R; specimen(s): Gwee, A.T. GAT 211.
Bruguiera gymnorrhiza (L.) Lam. ex Savigny; habit : tree; 7m tall; habitat : mangrove;
conservation status: R; specimen(s): Gwee, A.T. GAT 215.
Bruguiera parviflora (Roxb.) Wight & Arn. ex Griff.; habit : tree; occurrence: rare;
habitat: mangrove; conservation status: En; specimen(s): Gwee, A.T. GAT 329.
Carallia brachiata (Lour.) Merr.; habit : tree; habitat: coastal forest; conservation
status: R; specimen(s): Gwee, A.T. GAT 378.
Ceriops decandra (Griff.) W. Theob.; habit : tree; occurrence: frequent; habitat:
mangrove; specimen(s): Gwee, A.T. GAT 256.
Ceriops tagal (Pers.) C.B. Rob.; habit : tree; habitat: mangrove; conservation Status:
R; specimen(s): Gwee, A.T. GAT 261.
Rhizophora apiculata Blume; habit : tree; 5m tall; occurrence: frequent; habitat:
mangrove; conservation status: R; specimen(s): Gwee, A.T. GAT 240.
Rhizophora mucronata Lam.; habit : tree; 3m tall; occurrence: frequent; habitat:
mangrove; conservation status: R; specimen(s): Gwee, A.T. GAT 214.
Rhizophora stylosa Griff.; habit : tree; occurrence: rare; habitat : mangrove;
conservation status: V; specimen(s): Gwee, A.T. GAT 267.
Rubiaceae
Aidia auriculata (Wall.) Ridsdale; habit : climber; habitat : rocky shore; specimen(s):
Gwee, A.T. GAT 356.
Aidia densiflora (Wall.) Masam.; habit : tree; occurrence: frequent; habitat: coastal
forest; conservation status: V; specimen(s): Gwee, A.T. GAT 80.
Canthium confertum Korth.; habit : tree; 3m tall; habitat: coastal forest; conservation
status: R; specimen(s): Gwee, A.T. GAT 149.
Canthium horridum Blume; habit: shrub; habitat: coastal forest; conservation status:
R; specimen(s): Gwee, A.T. GAT 383.
304 Gard. Bull. Singapore 55 (2003)
Diplospora malaccensis Hook.f.; habit : tree; 3m tall; habitat: coastal forest;
conservation status: En; specimen(s): Gwee, A.T. GAT 83.
Guettarda speciosa L.; habit: tree; 6m tall; habitat: rocky shore; conservation status:
R; specimen(s): Gwee, A.T. GAT 163.
Gynochthodes coriacea Blume; habit : climber; habitat: wasteland; conservation
status: R; specimen(s): Gwee, A.T. GAT 200.
Ixora congesta Roxb.; habit : shrub; 2m tall; habitat: coastal forest; conservation
status: R; specimen(s): Leong, P. PL 1.
Ixora lobbii King & Gamble; habit : shrub; specimen(s): Gwee, A.T. GAT 293.
Lasianthus hirsutus (Roxb.) Merr.; habit : shrub; occurrence: rare; habitat: coastal
forest; conservation status: Ex; specimen(s): Leong, P. PL 9. (formally known as L.
cyanocarpus).
Morinda citrifolia L.; habit: tree; habitat : wasteland; conservation status: exotic;
specimen(s): Gwee, A.T. GAT 238.
Morinda umbellata L.; habit : climber; occurrence: frequent; habitat : coastal forest;
conservation status: common; specimen(s): Leong, P. PL 4.
Oxyceros fragrantissima (Ridl.) K.M. Wong; habit : climber; habitat: rocky shore;
conservation status: R; specimen(s): Gwee, A.T. GAT 357.
Oxyceros longiflora (Lam.) T. Yamaz.; habit : shrub; 1m tall; habitat: coastal forest;
conservation status: R; specimen(s): Gwee, A.T. GAT 136.
Psychotria malayana Jack, habit: shrub; habitat: coastal forest; conservation status:
R; specimen(s): Gwee, A.T. GAT 321.
Scyphiphora hydrophyllacea C.F. Gaertn.; habit : tree; 2m tall; habitat : sandy shore;
conservation status: R; specimen(s): Gwee, A.T. GAT 170.
Timonius flavescens (Jack) Baker; habit : tree; 1.5m tall; habitat: coastal forest;
conservation status: R; specimen(s): Gwee, A.T. GAT 125.
Survey of Chek Jawa, Singapore 305
Timonius wallichianus (Korth.) Valeton; habit : tree; habitat: coastal forest;
conservation status: R; specimen(s): Gwee, A.T. GAT 389.
Rutaceae
Clausena excavata Burm.f.; habit : shrub; occurrence: frequent; habitat : coastal
forest; conservation status: R; specimen(s): Gwee, A.T. GAT 97.
Glycosmis chlorosperma (Blume) Spreng. var. chlorosperma; habit : shrub; habitat :
coastal forest; conservation status: R; specimen(s): Gwee, A.T. GAT 87.
Melicope hookeri T.G. Hartley; habit : tree; habitat : coastal forest; conservation
status: R; specimen(s): Gwee, A.T. GAT 294.
Sapindaceae
Allophylus cobbe (L.) Raeusch.; habit : shrub; habitat : Sandy shore, wasteland;
conservation status: R; specimen(s): Gwee, A.T. GAT 265.
Guioa pubescens (Zoll. & Moritzi) Radlk.; habit : tree; habitat : coastal forest;
conservation status: R; specimen(s): Gwee, A.T. GAT 71.
Lepisanthes rubiginosa (Roxb.) Leenh.; habit: tree; occurrence: frequent; habitat:
coastal forest; specimen(s): Gwee, A.T. GAT 88.
Mischocarpus sundaicus Blume; habit : tree; habitat: coastal forest; conservation
status: R; specimen(s): Gwee, A.T. GAT 175.
Sapotaceae
Chrysophyllum roxburghii G. Don; habit : tree; habitat : coastal forest; specimen(s):
Leong, P. PL 19.
Pouteria linggensis (Burck) Baehni; habit : tree; habitat : rocky shore; specimen(s):
Gwee, A.T. GAT 107.
Pouteria malaccensis (C.B. Clarke) Baehni; habit : tree; habitat: coastal forest;
conservation Status: R; specimen(s): Gwee, A.T. GAT 379.
Pouteria obovata (R.Br.) Baehni; habit : tree; habitat: coastal forest; conservation
status: R; specimen(s): Gwee, A.T. GAT 73.
306 Gard. Bull. Singapore 55 (2003)
Scrophulariaceae
Scoparia dulcis L.; habit: herb; 0.75m tall; habitat: wasteland; conservation status:
exotic; specimen(s): Gwee, A.T. GAT 191.
Simaroubaceae
Brucea javanica (L.) Merr.; habit : tree; habitat : wasteland; conservation status: R:
specimen(s): Gwee, A.T. GAT 361.
Eurycoma longifolia Jack; habit: tree; habitat: coastal forest; conservation status: R:
specimen(s): Gwee, A.T. GAT 58.
Sonneratiaceae
Sonneratia alba J.J.Sm.; habit : tree; 4m tall; habitat : mangrove; specimen(s): Gwee,
A.T. GAT 100.
Sonneratia ovata Back.; habit: tree; conservation status: common; specimen(s): Gwee,
AT GAP 271.
Sterculiaceae
Heritiera littoralis Dryand.; habit: tree; habitat : mangrove; conservation status: R;
specimen(s): Gwee, A.T. GAT 255.
Scaphium macropodum (Miq.) Beumée ex Heyne; habit : tree; habitat : rocky shore;
conservation status: V; specimen(s): Gwee, A.T. GAT 178.
Symplocaceae
Symplocos fasciculata Zoll.; habit : tree; habitat: coastal forest; conservation status:
R; specimen(s): Gwee, A.T. GAT 221.
Thymelaeaceae
Linostoma pauciflorum Griff.; habit : climber; occurrence: frequent; habitat: coastal
forest; conservation status: R; specimen(s): Gwee, A.T. GAT 98.
Turneraceae
Turnera ulmifolia L.; habit: herb; 0.5m tall; habitat: wasteland; conservation status:
exotic; specimen(s): Gwee, A.T. GAT 13.
Ulmaceae
Trema tomentosa (Roxb.) Hara; habit: shrub; habitat: coastal forest; conservation
status: common; specimen(s): Leong, P. PL 6.
Survey of Chek Jawa, Singapore 307
Verbenaceae
Clerodendrum inerme (L.) Gaertn.; habit : shrub; occurrence: f srequent; habitat :
sandy shore; conservation status: R; specimen(s): Gwee, A.T. GAT 213.
Clerodendrum laevifolium Blume; habit : shrub; 2m tall; occurrence: frequent; habitat:
coastal forest; conservation status: R; specimen(s): Gwee, A.T. GAT 60.
Premna serratifolia L.; habit : shrub; 1.8m tall; habitat: sandy shore; conservation
status: R; specimen(s): Gwee, A.T. GAT 209.
Vitex pinnata L.; habit : tree; habitat: coastal forest; conservation status: common;
specimen(s): Gwee, A.T. GAT 101.
Vitaceae
Ampelocissus elegans (Kurz) Gagnep.; habit : climber; habitat: coastal forest;
conservation status: R; specimen(s): Gwee, A.T. GAT 311.
Cissus repens Lam.; habit: climber; habitat : wasteland; conservation status: R;
specimen(s): Leong, P. PL 15.
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Gardens’ Bulletin Singapore 55 (2003) 309-315
OBITUARY
Dr Chang Kiaw Lan, 31 July 1927-14 August 2003
Dr Chang Kiaw Lan was appointed Botanist to the Singapore Botanic Gardens Service
on 7 July 1959 and as Keeper of the Herbarium in 1970. She retired on 30 July 1987.
Throughout most of her career she was based at the Botanic Gardens at Cluny Road,
attached to and in charge of the Herbarium there, whence she made the acquaintance
of a great number of taxonomic botanists and other botanical visitors from throughout
the world. Her role in editing the Gardens’ Bulletin, Singapore for 17 years was
particularly significant because it was during a period when taxonomic research at
the Gardens was at an ebb and yet the Bulletin was the only actively produced,
dedicated botanical journal in Southeast Asia.
Kiaw Lan was born in Koetaradja (today called Banda Aceh) in north Sumatra,
to Mr Thio Kie San, Kapitan or leader of the Chinese community there, and Madam
Jong Lie Nio, when Indonesia was under Dutch administration. She was their second
daughter and youngest of six children (including an adopted first child). Kiaw Lan
attended early schooling there, which was interrupted by the Second World War, and
continued her education in Penang, Malaysia, from January 1948 until July 1953. (It
was in Penang that she began to use the name Chang Kiaw Lan consistently, one of
several translated phonetic versions of her name in Chinese that included Thio Kiauw
Lan, Tjong Kiauw Lian and Tjong Su Yun!) She moved to Singapore to read Botany
and Philosophy at the University of Malaya (1953-1958) and in 1959 gained an Upper
Il Honours in Botany. She obtained a Certificate of Naturalisation under the British
Nationality Act to stay in Singapore in May 1959 just prior to her joining the Botanic
Gardens Service and in December, 1959, became a Singapore citizen. Her first field
trip as a botanist in the Botanic Gardens appears to have been during 21—28 September
1959, in the Sedili area in Johore, in the company of Mohamad Shah, a plant collector
who eventually rose to become Herbarium Assistant and Research Officer and who
would ably assist her with the functions of the Herbarium throughout her career there.
Following her appointment to the Botanic Gardens, under H.M. Burkill’s
directorship, she was awarded a British Commonwealth Scholarship from October
1962 to July 1964 for a PhD in Botany at the University of Cambridge. She worked
under the supervision of E.J.H. Corner, Professor of Tropical Botany at the Botany
School, an exceedingly talented botanist with wide interests covering mycology and
seed plants. Corner had served as an Assistant Director at the Singapore Botanic
Gardens and was already well known as the author of Wayside Trees of Malaya by the
time the Second World War overtook events in Singapore in 1942. Kiaw Lan was
attached to Newnham College and completed her thesis, “The Structure and Taxonomy
of Lentinus Fr.” (xii + 247 p., 65 pl.) in July 1965. The thesis provided a resume of
310 Gard. Bull. Singapore 55 (2003)
Chang Kiaw Lan at the Singapore Herbarium upon return from Cambridge, 1965 (A); at her
desk, December 1974 (B); attending a friend’s reception, sometime 1970s (C); and while on
tour in Vietnam shortly after retirement (D). (Courtesy of Christina Tan)
characters of taxonomic value in the classification of the gill-fungus Lentinus, which
is well-represented in Malaysia and Singapore, and provided a new definition and
typification for the genus, previously poorly circumscribed. It also confirmed the
value of hyphal analysis as a taxonomic method. Then, accurate identification of the
Asian tropical species was virtually impossible because there were no keys.
She resumed her service at the Gardens on 21 September 1965, coming back
to arather active schedule trying to settle in, but based at a temporary work station in
the library while the office she was to occupy in the new three-floor Herbarium
(completed in 1964) was still used by James Sinclair. He had been Curator since
Obituary: Chang Kiaw Lan 311
1948, retired in 1963, was re-employed until 1965, when he worked in an honorary
capacity until 1967. In August/September 1966 she attended the Eleventh Pacific
Science Congress in Tokyo, where she presented a paper on “Criteria for the
classification of Lentinus species”. There, she also attended the Marine Park
Symposium and the 1“ U.S.-Japanese Algal Culture Conference in Ueno Park. Her
trip was sponsored by the Singapore Industrial Research Unit and financed by the
American Embassy and the Asia Foundation. During 1-5 October in the same year,
she collected fungi in the Cameron Highlands, Peninsular Malaysia, together with
Professor Corner and Dr Chew Wee Lek, the first local graduate appointed as botanist
in 1956, obtaining his PhD from Cambridge in 1965.
In 1967, the Botanic Gardens was about to enter a trying period. Following
her training in mycology, Kiaw Lan had already identified the need to carry out
developmental studies of basidiomycete fungi because of the paucity of any
information on the tropical species. For this, she realized that facilities for experimental
work at the Gardens had to be set up. On the other hand, work on orchid culture
(which involved another botanist, Hardial Singh, engaged in 1963) was already well
established. That year, when the Singapore government’s beautification programme
became significant, she was also asked to be in charge of advisory work for the new
Tree Planting Campaign.
Then, on 2 May 1968, she was seconded to the Primary Production
Department, engaged in mushroom cultivation research. Kiaw Lan’s duties in 1968
included studying the pests and diseases of cultivated plants, investigating the
commercial production of edible mushrooms, and mycological research and routine
maintenance of fungal collections. She was particularly interested in the mushroom
genus Jermitomyces, noting, “It is well-known that the termite mushroom is the most
delicious in tropical countries.” The name of the mushroom alludes to its association
with the subterranean nest of Odontotermes termites, which do not build conspicuous
mounds. The fungus is cultivated in so-called fungus gardens in the nest chambers of
these termites, and helps in the digestion of cellulosic material brought in by the
termites. But just over a year later, on 1 July 1969, she was returned after Dr Chew
drew attention to an “acute shortage of botanists in the Botanic Gardens”. In 1970,
Chew assumed the duties of Director and Kiaw Lan became the Keeper of the
Herbarium. So her duties continued to be defined around mycology but the contexts
were different and rather wide-ranging.
Ironically, her role became more significant as taxonomic research waned in
Singapore, particularly after the retirement of Burkill and departure of Chew, just
before the Garden came under the organisation of The Parks & Recreation Service in
1973. The new department was overwhelmingly dedicated to the development of a
superior Garden City that has come to be Singapore’s signature theme. In 1970, Geh
Siew Yin was appointed botanist, but she was moved to administrative duties as
Assistant Commissioner of the Garden in 1973. By 1975, Hardial Singh was also
312 Gard. Bull. Singapore 55 (2003)
transferred to non-research duties in the expanding Parks & Recreation Department.
Hence, while one important aspect was fulfilled, resulting in a very successful
urban and suburban horticultural management, the other, which would have been
based on a traditional strength in tropical botanical research, faded. In retrospect,
both areas could have flourished, as Singapore could well afford such development
and was central, in terms of service and expertise, to a rapidly developing resource-
rich region. Whereas, previously, many standard works on the botany of the region,
such as Wayside Trees of Malaya, Malayan Wild Flowers, and numerous taxonomic
revisions in the Gardens’ Bulletin, were crafted by botanists at the Singapore Botanic
Gardens, the 1970s and 1980s saw a dearth of such activity. In that period, the
taxonomic output from the Gardens itself declined sharply and the key regional work
of the time, the four-volume Tree Flora of Malaya, a project spanning over twenty
years and involving 19 botanists, had no Botanic Garden participation in authorship.
(It was to be in the late 1980s before the resuscitation process for the taxonomic and
floristic research functions in the Garden began gathering momentum, leading to
renewed botanical activity so evident since the 1990s.)
In her 1948 Pupil’s Report Book at the Convent School, Penang, where Kiaw
Lan first came to study from Sumatra, she was said to have “faulty accents” and it
was recommended she paid more attention to reading and pronunciation. From this
uncertain start, she made great progress in mastering English until, later, even members
of the English Department at the University of Malaya admired her command of the
language (Zaibun Siraj, pers. comm.). She was outgoing and found all manner of
expression interesting. In A Private Landscape, a 1967 anthology of contemporary
Malaysian poetry by David Ormerod, who taught in the English Department at the
University of Malaya, Kiaw Lan contributed a poem called “The woman who cried
Adohi’”’, a tongue-in-cheek poke at male perception, under the pen-name Muh Lan.
Kiaw Lan spoke English, Dutch, French, Chinese (Mandarin and the Hokkien and
Hakka dialects), Indonesian Malay and some German. If there was a single area that
was Kiaw Lan’s forte, it would have been language, and this was much recognized as
she became involved with the editing of the Gardens’ Bulletin, Singapore, the Botanic
Gardens’ official journal.
Ranu Dally, one of her best friends from early youth, recalls that they
appreciated the Malay/Indonesian cultures by virtue of their background (Ranu from
Kelantan, Kiaw Lan from Sumatra). Kiaw Lan was much fascinated by things
Indonesian and was always captivated by interesting hand-made textiles, including
especially cloths with bold patterns (called ikat), which she collected, and other objects
of art from the region. In this and in Dutch-Indonesian adaptations and habits, she
found much common ground with another special friend, Singapore philantropist
Ted di Ponti. She had also learnt pottery under John Shelly in Devon in August 1963,
close-up photography with the Nature Society in 1978, and newspaper design and
layout in a course run by the School of Journalism of the Times Press Foundation in
Obituary: Chang Kiaw Lan 513
December 1984.
Kiaw Lan took charge of the Herbarium, a vast specimen archive of the
botanical wealth of the Malaysian-Indonesian region, and the Gardens’ botanical library
for a long time, when development opportunities for these were scarce. She was a
prolific correspondent and prompt in addressing requests from visitors to the Gardens
and Herbarium. She was kept busy with the Gardens’ Bulletin, and was careful,
judicious and accurate in her editorial work and correspondence, which became one
of its hallmarks. Later, looking back, she was especially proud of several works relating
to a former supervisor she admired greatly. Together with David Mabberley, Kiaw
Lan had also edited the Corner Festschrift (Tropical Botany, Essays presented to EJH
Corner for his Seventieth Birthday, Gardens’ Bulletin Vol. 29), published in 1977.
She helped bring out Corner’s The Freshwater Swamp-forest of South Johore and
Singapore, which was issued as the Bulletin’s Supplement No. 1 (1978). She also
assisted in the preparation of other works, such as the much applauded Proceedings
of the 11" World Orchid Conference held in Singapore (published in 1985). After
1987, the Bulletin came under the editorship of Geh Siew Yin, with whom Kiaw Lan
had worked for some years prior to retirement. Geh also assumed Keepership of the
Singapore Herbarium from 1987, amid other duties.
Outside the Gardens, Kiaw Lan was treasurer or committee member of the
Singapore Branch of the Malayan Nature Society from 1970 to 1980, including being
Branch Chairman for 1976-77. She served on the committee of the Alliance Francaise
de Singapour as member or treasurer between 1971 and 1980, and was one of its
registered School Managers. In 1976, she was honoured by the French government
as a Knight in the National Order for Academic Excellence. She was also a member
of the Amateur Radio Society and the University of Singapore Society. Kiaw Lan
was also involved in the Nature Reserves Board as secretary for a number of years.
After she retired at age 60, Kiaw Lan tried various editing assignments,
including technical writing and editing with a company that printed user manuals for
radio equipment, and editing the odd book and magazine issue outside Botany. She
learnt various word-processing software programmes on computer and even word-
processed some books for the Braille Production Unit of the Visually Handicapped
Society. She took more interest in the apartment complex where she stayed, as a
member of its management council, found occasion for treating friends to her cooking,
but also went on tours from Singapore. Old botanical acquaintances continued to call
on her, and she enjoyed accommodating one or two as they visited the Herbarium for
work. She continued to entertain her visitors at the Cricket Club, which she often
used as a member.
Several jobs gave her a sense of familiarity. Kiaw Lan was sought for indexing
the 3" edition of Corner’s Wayside Trees of Malaya, which was published in 1988 by
the Malayan Nature Society. For such a classic work, it was unfortunate that the
cover picture on Volume | was captioned with the wrong name (Alstonia angustifolia
314 Gard. Bull. Singapore 55 (2003)
instead of Alstonia angustiloba) during production. When Kiaw Lan wrote a personal
note to explain the mistake, a very magnanimous professor wrote back to one of his
favourite former students: “J never supposed that the mistake. . .was your doing. .
.Perhaps the photographer made the mistake. Anyhow it is a battered old tree that
has lost its top and not a picture that I would have chosen. However, old trees must
not be scorned. . .I have always been proud of the book and Ed. 3 is your gift and
more to S.E.Asia.” Later Corner sent another manuscript, Moments Botanique, a
biography of botanical tales from the Malay Peninsula to the Solomon Islands and
South America, for Kiaw Lan to word-process from handwritten material, as he
believed she could interpret his handwriting well enough and she offered good
comment and suggestion. Another job was the compilation of the master index
accompanying the issue of the final (fourth) volume of the Tree Flora of Malaya,
published in 1989 by Longman.
It was to be in the 1990s before positions in taxonomic botany became
available again in the Singapore Botanic Gardens. It was during this period, after the
departure of Burkill and Chew, and when the new effort for the Garden City ironically
overshadowed other aspects at the Botanic Gardens, that Chang Kiaw Lan remained
steadfast in her role as the Keeper of the Singapore Herbarium and a fixture in the
affairs of the Gardens’ Bulletin. Today, both herbarium and journal continue their
development, having emerged from the very edge of botanical history.
Kiaw Lan suffered a minor heart attack on 17 February 1987 and was
diagnosed with cancer in April 2002, when also a condition of dementia became
more obvious, with recurring memory lapses. She was indeed most fortunate to have
had a special friend in Christina Tan, who with her husband Christopher Lim
accommodated Kiaw Lan into their lives, and later arranged for her consistent care in
July 2002. Kiaw Lan was admitted to Hospice care in July 2003, and died on 14
August 2003. Kiaw Lan leaves behind a brother, Paul Gaudiamo (Indonesia), and a
sister, Meta de Visser (U.S.A.), nephews, nieces, and their families; her other siblings
have passed on. In her will, she left her body to medical research and also wished for
a gathering of her friends to be arranged. Many of her family members and friends
were present at the Memorial Gathering for her on 19 August 2003 at the Orchard
Guild House on Grange Road, Singapore. The venue was chosen by Christina to
recall Kiaw Lan’s liking for meeting with friends at the original Guild House of the
National University of Singapore Society at Evans Road, not far from the Botanic
Gardens.
Does not, then, the charm of life remain for me long and lasting,
when my burdens are gone, my surroundings are quiet, my body is
free and my heart leisurely?
Obituary: Chang Kiaw Lan
>
J Lin Yutang, “The Importance of Living’
K.M. Wong
University of Malaya,
Kuala Lumpur
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